Does vicariance shape biotas? Biogeographical tests of the vicariance model in the north‐west European land snail fauna

Aim To investigate the importance of vicariance in shaping the north‐west European land snail fauna. Location North‐west Europe. Methods We tested whether there is a non‐random congruence, i.e. a clustering of species ranges, using a Monte Carlo procedure with a null model that generates range data sets such that their range size distribution, the species richness distribution of the geographic cells and the spatial autocorrelation of the occurrences of a taxon approach the parameters in the real data set. Biotic elements, groups of species with similar ranges, were delimited with Model based Gaussian clustering. The prediction that closely related species belong to different biotic elements, has been tested with a chi‐square test. Results The distribution areas of the north‐west European land snail species are significantly clustered as predicted by the vicariance model. One widespread and seven regional biotic elements were identified. Contrary to the predictions of the vicariance model, closely related north‐west European land snail species belong significantly more often to the same biotic element than should be expected by chance. Main conclusions The clustering of closely related north‐west European land snail species within the same biotic element indicates that speciation modes other than vicariance were frequent or that the imprint of vicariance on the ranges was obscured by extensive post‐speciational dispersal. Extensive dispersal may have been caused by Pleistocene climatic fluctuations. The core areas of the regionally restricted biotic elements might indicate the positions of glacial refugia of land snails.     

Beringia as a high-latitude engine of avian speciation

Beringia is a biogeographically dynamic region that extends from northeastern Asia into northwestern North America. This region has affected avian divergence and speciation in three important ways: (i) by serving as a route for intercontinental colonisation between Asia and the Americas; (ii) by cyclically splitting (and often reuniting) populations, subspecies, and species between these continents; and (iii) by providing isolated refugia through glacial cycles. The effects of these processes can be seen in taxonomic splits of shallow to increasing depths and in the presence of regional endemics. We review the taxa involved in the latter two processes (splitting–reuniting and isolation), with a focus on three research topics: avian diversity, time estimates of the generation of that diversity, and the regions within Beringia that might have been especially important. We find that these processes have generated substantial amounts of avian diversity, including 49 pairs of avian subspecies or species whose breeding distributions largely replace one another across the divide between the Old World and the New World in Beringia, and 103 avian species and subspecies endemic to this region. Among endemics, about one in three is recognised as a full biological species. Endemic taxa in the orders Charadriiformes (shorebirds, alcids, gulls, and terns) and Passeriformes (perching birds) are particularly well represented, although they show very different levels of diversity through evolutionary time. Endemic Beringian Charadriiformes have a 1.31:1 ratio of species to subspecies. In Passeriformes, endemic taxa have a 0.09:1 species-to-subspecies ratio, suggesting that passerine (and thus terrestrial) endemism might be more prone to long-term extinction in this region, although such ‘losses’ could occur through their being reconnected with wider continental populations during favourable climatic cycles (e.g. subspecies reintegration with other populations). Genetic evidence suggests that most Beringian avian taxa originated over the past 3 million years, confirming the importance of Quaternary processes. There seems to be no obvious clustering in their formation through time, although there might be temporal gaps with lower rates of diversity generation. For at least 62 species, taxonomically undifferentiated populations occupy this region, providing ample potential for future evolutionary diversification.     

Genetic differentiation in the Trinidad endemic Mannophryne trinitatis (Anura: Aromobatidae): Miocene vicariance,in situ diversification and lack of geographical structuring across the island

Trinidad offers a unique study system within the Caribbean to assess the processes and patterns of amphibian speciation. We used mitochondrial DNA sequences to investigate the phylogenetic relationships and patterns of intraspecific genetic variation of Mannophryne trinitatis from Trinidad. Molecular clock estimates point to a genetic split between M. trinitatis and its sister species, M. venezuelensis, dating to the Late Miocene (c. 7–8 Mya), suggesting vicariance as a means of speciation when Trinidad pulled apart from northern Venezuela. M. trinitatis phylogenetic population analyses from ten Northern Range and four Central Range localities recovered three well‐resolved clades: a larger clade formed by haplotypes from Northern Range localities and two additional clades, one formed by haplotypes from the Central Range and another including haplotypes from Northern Range localities and one haplotype from the Central Range. Overall, our results show that the genetic diversity in M. trinitatis is not geographically structured but it is distributed among the various Northern and Central Range localities. In congruence with the vicariance speciation hypothesis, we attribute M. trinitatis present distribution and lack of genetic structure to multiple admixture events caused by climate changes that severely affected the topology of Trinidad throughout the Pliocene/Pleistocene periods.     

Phylogenetics, zoogeography, and the role of dispersal and vicariance in the evolution of the Rana catesbeiana (Anura: Ranidae) species group

The Rana catesbeiana species group consists of seven species, each variously distributed across eastern North America. We estimated the evolutionary relationships among 31 exemplars and used the phylogenetic hypothesis to examine the potential modes of speciation and relative role of dispersal in the evolution and zoogeography of this species group. Phylogenetic relationships based on 1554 combined base pairs of the cytochrome b and ND2 mitochondrial genes suggest that the species are closely related, having undergone rapid radiation from a common ancestor during the late Miocene or Pliocene. A Pleistocene origin for the rare R. okaloosae is suggested by its pattern of paraphyly with R. clamitans and by its geographically restricted distribution, although hybridization as the explanation for paraphyly cannot be ruled out. Dispersal–vicariance analysis suggested a Coastal Plain biogeographical region origin of the species group, supporting the notion that the region was an important centre of anuran diversification, with post-speciation dispersal playing a major role in explaining the distribution of the widespread species, R. catesbeiana , R. clamitans, and R. septentrionalis . High sea levels during the late Tertiary, greatly reducing and insularizing parts of the southern Coastal Plain region may have played a major role in the diversification of this group.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 601–624.     

Biostratigraphy and sedimentary evolution of Late Miocene and Pliocene continental deposits of the Granada Basin (southern Spain)

The Late Miocene and Pliocene continental sediments in the Granada Basin (southern Spain) have yielded large amounts of fossil small mammals in 37 localities from 11 sections. The aim of this paper is to integrate faunistic, stratigraphic, and sedimentary criteria to unravel the geological history of the continental infilling of the basin. The palaeontological study has led to a detailed biozonation on the basis of rodents, which helps to correlate in detail the different sedimentary units found in the basin, and to follow the changes of the different sedimentary systems and their palaeogeographical evolution through time. Combination of the proposed biostratigraphy and the reinterpretation of the magnetostratigraphic analyses of the Barranco del Purcal section allows us to assign an absolute age slightly older than 5.23 Ma to the Turolian–Ruscinian boundary (MN13‐MN14).     

Pliocene forest dynamics as a primary driver of African bird speciation

Aim Montane tropics are areas of high endemism, and mechanisms driving this endemism have been receiving increasing attention at a global scale. A general trend is that climatic factors do not explain the species richness of species with small to medium‐sized geographic ranges, suggesting that geological and evolutionary processes must be considered. On the African continent, several hypotheses including both refugial and geographic uplift models have been advanced to explain avian speciation and diversity in the lowland forest and montane regions of central and eastern Africa; montane regions in particular are recognized as hotspots of vertebrate endemism. Here, we examine the possible role of these models in driving speciation in a clade of African forest robins. Location Africa. Methods We constructed the first robustly supported molecular phylogenetic hypothesis of forest robins. On this phylogeny, we reconstructed habitat‐based distributions and geographic distributions relative to the Albertine Rift. We also estimated the timing of lineage divergences via a molecular clock. Results Robust estimates of phylogenetic relationships and clock‐based divergences reject Miocene tectonic uplift and Pleistocene forest refugia as primary drivers of speciation in forest robins. Instead, our data suggest that most forest robin speciation took place in the Late Pliocene, from 3.2 to 2.2 Ma. Distributional patterns are complex, with the Albertine Rift region serving as a general east–west break across the group. Montane distributions are inferred to have evolved four times. Main conclusions Phylogenetic divergence dates coincide with a single period of lowland forest retraction in the late Pliocene, suggesting that most montane speciation resulted from the rapid isolation of populations in montane areas, rather than montane areas themselves being drivers of speciation. This conclusion provides additional evidence that Pliocene climate change was a major driver of speciation in broadly distributed African animal lineages. We further show that lowland forest robins are no older than their montane relatives, suggesting that lowland areas are not museums which house ‘ancient’ taxa; rather, for forest robins, montane areas should be viewed as living museums of a late Pliocene diversification event. A forest refugial pattern is operating in Africa, but it is not constrained to the Pleistocene.     

Late Miocene vegetation and climate reconstruction based on pollen data from the Sofia Basin (West Bulgaria)

The analysis of fossil palynomorh assemblages in the Late Miocene freshwater sediments of the Sofia Basin (West Bulgaria) was done to collect data on the vegetation and climate dynamics during the Late Miocene. On the basis of pollen data, we described the main palaeocomunities developed in the region. The mixed mesophytic forests dominated the vegetation in which species of Quercus, Ulmus, Zelkova, Fagus, Carpinus, Betula, Castanea, Corylus, Pterocarya, Carya, Juglans, and Eucommia played important roles. Swamp forests were also recorded, including Taxodiaceae, Alnus, Glyptostrobus, Nyssa, and Myrica. Herbaceous vegetation was distributed in the middle part of the section, with a maximum of 35.5%. The vegetation dynamic passes through several phases, which were associated with changes in paleoclimate and palaeoecological conditions. Coexistence Approach (CA) was applied to palynological data to calculate four climatic parameters. The values of coexistence intervals for mean annual temperatures are 13.6–16.6 °C, with winter temperatures being 3.7–6.6 °C and summer temperatures being 23.6–27.8 °C. Mean annual precipitation ranged most frequently between 828 and 1308 mm. The palaeoclimatic reconstruction illustrates existence of a warm-temperate and relatively humid climate with higher mean annual temperature than the present day climate.     

Euroceros bulgaricus gen. nov., sp. nov. from Hadzhidimovo (SW Bulgaria) (Late Miocene) - the first European record of Hornbills (Aves: Coraciiformes)

A new genus and species of ground-hornbill are described from the Late Tertiary (Late Miocene) fluviative deposits in Hadzhidimovo (Hadzhidimovo-1, or Hadzhidimovo-Girizite locality), in southwestern Bulgaria. The cranial half of a scapula and the distal third of a femur have been recovered. The morphology distinguishes the specimen from the recent species of Bucorvus, the closest genus. Most interesting is the morphology of the scapula (the holotype), which differs considerably from the ground-hornbills in some structures (e.g. the clavicular joint) while being surprisingly similar in other structures (e.g. the humeral articular joint). The last peculiarity may indicate that the fossil species was a more primitive form having relatively poorer flying capacity in comparison to other ground-hornbills. This first record of Bucerotidae in Europe proves the occurrence of the family in the Tertiary of the western parts of the Eurasian continent. It came from the richest Bulgarian locality of the Hipparion fauna, from which ca. 36 species of large mammals and birds are now known.     

Speciation and radiations track climate transitions since the Miocene Climatic Optimum: a case study of southern African chameleons

Aim The high amount of species diversity concentrated in southern Africa has been attributed to palaeoclimatic factors, and the timing of radiations in some taxa corresponds to global palaeoclimatic trends. Using dwarf chameleons (Bradypodion: Chamaeleonidae) as a model system, we explored the relationship between palaeoclimatic fluctuations and cladogenesis with respect to both temporal and spatial patterns in an effort to understand the process of speciation in southern Africa. Location South Africa, with particular emphasis on the Cape Floristic Region and the Maputaland–Pondoland–Albany hotspot. Methods Mitochondrial sequence data (ND2 and 16S) were used to estimate the timing of major radiations and to examine the number of lineages through time. A dated phylogeny was constructed using Bayesian phylogenetic reconstruction, and a Bayesian relaxed molecular clock was used to estimate divergence times. Spatial data and lineage‐through‐time plots were used to identify geographic regions that underwent diversification in connection with major climatic events. Both parsimony and likelihood optimizations of habitat type on the phylogeny were used to determine whether major habitat shifts have occurred. On a coarse scale (half‐degree grid cells), phylogenetic diversity (sum of the branch lengths linking terminals) was compared with species richness (absolute number of species) to identify areas of conservation importance. Results The complete species phylogeny of dwarf chameleons shows that the timing and mode of diversification exhibit spatio‐temporal patterns that link to phases in the evolution of southern Africa’s climate over the last 14 Myr. Optimizations of habitat on the phylogenetic tree show a progression from closed to open habitats since the Mid‐Miocene, corresponding to the shift from C₃ to C₄ environments, and later with the development of south‐western Africa’s winter‐rainfall regime. These shifts are not simultaneous across the region, with different geographic centres of diversity generated during different time periods. Main conclusions Regions that are prominent centres of chameleon diversification are encompassed by the current biodiversity hotspots as shown by chameleon species richness and phylogenetic diversity. Diversity within the Cape Floristic Region appears to be the result of a Late Pliocene radiation, whereas the diversity encompassed within the Maputaland–Pondoland–Albany hotspot is an aggregate of asynchronous radiation events, probably influenced by lineage losses. Overall, dwarf chameleons have experienced a shift in habitat types, with recent radiations occupying open habitats, and older lineages persisting in relictual forested habitats, corresponding to the continental shift of vegetation types since the Miocene Climatic Optimum.     

Late Miocene lagomorphs from the Republic of Moldova

The aim of this paper is to enlarge knowledge about still poorly documented and understood record of Lagomorpha (Mammalia) from the Late Miocene of Moldova. The lagomorph material from two new Late Miocene localities, Razeni (MN11/12), Gradishte (MN12), and re-sampling locality Chimislyia (MN12), of Southern Moldova, is described in detail here. The localities yielded small vertebrate fauna including five lagomorph taxa: Prolagus cf. oeningensis, “Proochotona” cf. eximia, Alilepus laskarewi, “Proochotona” sp., and Alilepus sp. P. cf. oeningensis is described for the first time in the studied area. A short review of the lagomorph record from the Republic of Moldova is provided. This contribution outlines the importance of the lagomorphs for biostratigraphic purposes, and highlighs the gaps in our knowledge related to the faunal succession in this area.     

Diversification within glacial refugia: tempo and mode of evolution of the polytypic fish Barbus sclateri

A diversity of evolutionary processes can be responsible for generating and maintaining biodiversity. Molecular markers were used to investigate the influence of Plio-Pleistocene climatic oscillations on the evolutionary history of taxa restricted to the freshwaters of a classical glacial refugium. Population genetic, phylogenetic and phylogeographical methods allowed the inference of temporal dynamics of cladogenesis and processes shaping present-day genetic constitution of Barbus sclateri , a polytypic taxon found in several independent river drainages in southern Iberian Peninsula. Results from different analyses consistently indicate several range expansions, high levels of allopatric fragmentation, and admixture following secondary contacts throughout its evolutionary history. Using a Bayesian demographical coalescent model on mitochondrial DNA sequences calibrated with fossil evidence, all cladogenetic events within B. sclateri are inferred to have occurred during the Pleistocene and were probably driven by environmental factors. Our results suggest that glaciation cycles did not inhibit cladogenesis and probably interacted with regional geomorphology to promote diversification. We conclude that this polytypic taxon is a species complex that recently diversified in allopatry, and that Pleistocene glaciation–deglaciation cycles probably contributed to the generation of biological diversity in a classical glacial refugium with high endemicity.     

Egarapithecus narcisoi, a new genus of Pliopithecidae (primates, catarrhini) from the Late Miocene of Spain

Pliopithecid remains from the Spanish locality of Torrent de Febulines (Late Vallesian, MN 10), consisting of right and left mandibular fragments with partial tooth rows and an isolated P(3) probably belonging to the same individual, are described and assigned to Egarapithecus narcisoi gen. et sp. nov. (Pliopithecidae, Crouzeliinae). This is a highly derived species dated at around 9 Ma (Ma = 10(6) years), representing the latest appearance of the family in the European continent. Morphologically it is the most distant member from the inferred primitive pliopithecid morphotype, displaying many autapomorphies that notably accentuate those of the remaining Crouzeliinae. A cladistic analysis based on lower cheek teeth, performed in order to tentatively assess the phylogenetic relationships of Egarapithecus within the Crouzeliinae, indicates that several equally parsimonious cladograms are possible in the light of current evidence. This is due to uncertainties regarding the position of Plesiopliopithecus and Crouzelia (here considered distinct genera), as a result of missing characters and the significant degree of homoplasy apparently involved in crouzeliine dental evolution. Whether Egarapithecus is more closely related to Crouzelia or to Anapithecus (the latter hypothesis tentatively favored here) cannot be definitively resolved with the currently available material and deserves further investigation. It is clear, however, that Egarapithecus is one of the more derived and specialized members of the Pliopithecidae.     

Genetic and biogeographic relationships of the racer goby Neogobius gymnotrachelus (Gobiidae: Teleostei) from introduced and native Eurasian locations

The racer goby Neogobius gymnotrachelus , along with several other neogobiin fishes, has been spreading north and west from its native Ponto-Caspian range for the past two decades via shipping and canals. It has been predicted as a likely future invader of the North American Great Lakes, where it would join its neogobiin relatives – the round and freshwater tubenose gobies. The present study is the first to analyse the population genetic and biogeographic relationships of the racer goby, establishing a baseline to aid interpretation of its future spread patterns and likely donor-recipient population relationships. The mitochondrial DNA cytochrome b gene was sequenced from representative areas of the racer goby's range, including rivers of the northern Black Sea and areas of spread upstream in the Danube River and outside the Ponto-Caspian region to the Vistula River in Poland. Results discerned nine haplotypes, with few shared among drainages of native rivers and most sites housing unique alleles. Racer goby populations significantly diverged among watersheds, supporting historic low migration and little non-anthropogenic gene flow. The Dnieper River was identified as a likely donor source for the Vistula River colonization, where appreciable variability suggests a relatively large number of founding genotypes.     

Biogeographical tests of the vicariance model in Mediterranean land snails

Aim To investigate the importance of vicariance in shaping land snail faunas. Location Three data sets of Mediterranean land snails were analysed: Helicoidea of the Iberian Peninsula and the complete land snail faunas of the central and eastern Aegean Islands and of Israel and Palestine. Methods The vicariance model predicts a clustering of species ranges. We tested for clustering of species ranges with a Monte Carlo simulation. For this simulation we used a null model that generates range data sets in such a way that their range size distribution, their species richness distribution and the spatial autocorrelation of the ranges approximate the parameters in the real data set. Biotic elements (clusters of species ranges) were delimited with model‐based Gaussian clustering. A second prediction of the vicariance model is that closely related species belong to different biotic elements. This was tested with a chi‐squared test. Results The distribution areas of the Iberian Helicoidea and the Israeli/Palestinian land snails are significantly clustered. The same is true for Israeli/Palestinian land snail species belonging to groups with geographically restricted species. However, the clustering is not significant in the complete central and eastern Aegean land snail data set, the Iberian Helicoidea species, and the central and eastern Aegean land snails belonging to groups with geographically restricted species. Contrary to the prediction of the vicariance model, closely related Iberian Helicoidea species and Israeli/Palestinian land snails belong significantly more often to the same biotic element than expected by chance. The null hypothesis that closely related species are homogeneously distributed across biotic elements cannot be rejected only for the data set that includes the Israeli/Palestinian land snails belonging to groups with geographically restricted species. Main conclusions The patterns found in the central and eastern Aegean land snail and the Iberian Helicoidea data sets do not correspond with the predictions of the vicariance model. This indicates that speciation modes other than vicariance were frequent, or that the distribution areas of many species in these faunas were largely modified by extensive post‐speciation dispersal and/or regional extinction. Hardly any possible vicariance events that might have contributed to the origin of the observed biotic elements in the Israeli/Palestinian land snail fauna could be identified. On the contrary, the biotic elements of the Israeli/Palestinian land snail fauna correspond with ecological factors. Vicariance has had only a limited influence on current biogeographical patterns in Mediterranean land snails.     

Cycles of trans‐Arctic dispersal and vicariance,and diversification of the amphi‐boreal marine fauna

The amphi‐boreal faunal element comprises closely related species and conspecific populations with vicarious distributions in the North Atlantic and North Pacific basins. It originated from an initial trans‐Arctic dispersal in the Pliocene after the first opening of the Bering Strait, and subsequent inter‐oceanic vicariance through the Pleistocene when the passage through the Arctic was severed by glaciations and low sea levels. Opportunities for further trans‐Arctic dispersal have risen at times, however, and molecular data now expose more complex patterns of inter‐oceanic affinities and dispersal histories. For a general view on the trans‐Arctic dynamics and of the roles of potential dispersal–vicariance cycles in generating systematic diversity, we produced new phylogeographic data sets for amphi‐boreal taxa in 21 genera of invertebrates and vertebrates, and combined them with similar published data sets of mitochondrial coding gene variation, adding up to 89 inter‐oceanic comparisons involving molluscs, crustaceans, echinoderms, polychaetes, fishes and mammals. Only 39% of the cases correspond to a simple history of Pliocene divergence; in most taxonomical groups, the range of divergence estimates implies connections through the entire Pliocene–Pleistocene–Holocene time frame. Repeated inter‐oceanic exchange was inferred for 23 taxa, and the latest connection was usually post‐glacial. Such repeated invasions have usually led to secondary contacts and occasionally to widespread hybridization between the different invasion waves. Late‐ or post‐glacial exchange was inferred in 46% of the taxa, stressing the importance of the relatively recent invasions to the current diversity in the North Atlantic. Individual taxa also showed complex idiosyncratic patterns and histories, and several instances of cryptic speciation were recognized. In contrast to a simple inter‐oceanic vicariance scenario underlying amphi‐boreal speciation, the data expose complex patterns of reinvasion and reticulation that complicate the interpretation of taxon boundaries in the region.     

Comparative osteology of the penguin‐like mid‐Cenozoic Plotopteridae and the earliest true fossil penguins,with comments on the origins of wing‐propelled diving

We compared the osteology of the late Eocene to early Miocene penguin‐like Plotopteridae from the North Pacific Basin with that of Paleocene stem group representatives of the Sphenisciformes and identified previously unrecognized similarities and differences. New data on the osteology of plotopterids, like the shape of the caudal end of the mandible, support a position of plotopterids outside the Suloidea, the clade formed by Sulidae, Phalacrocoracidae, and Anhingidae. However, as assumed by previous authors, the diving adaptations of plotopterids and sphenisciforms are likely to have evolved independently, and the resemblances in different parts of the postcranial skeleton therefore constitute one of the more striking examples of parallelism among tetrapods. We note that close relatives of both plotopterids and penguins forage by plunge diving. Whereas underwater locomotion of diving birds with a swimming ancestor is usually driven by the feet, we hypothesize that plotopterids and other wing‐propelled divers are more likely to have had volant ancestors that initiated diving by shallow plunges into the sea.     

Systematic revision of the Late Miocene sabre‐toothed felid Paramachaerodus in Spain

Abstract: A systematic revision of the sabre‐toothed cat genus Paramachaerodus  Pilgrim, 1913 is presented. Two species are recognized within Paramachaerodus, Pa. orientalis, and Pa. maximiliani, and the genus Promegantereon  Kretzoi, 1938 is retrieved to include Promegantereon ogygia. Material from the Turolian Spanish localities of Crevillente‐2 (MN 11, Alicante) and Las Casiones (MN 13, Teruel), which was previously assigned to Paramachaerodus, is now included in the tribe Metailurini. The exceptional discoveries at the Spanish Vallesian (MN 10, Madrid) fossil site of Batallones‐1 have made it possible to characterize the dentition and cranial anatomy of a previously very poorly known machairodontine cat, formerly included in Paramachaerodus as Pa. ogygia, which now can be distinguished from Pa. orientalis and Pa. maximiliani by the following features: canines without crenulations, P3 with a marked disto‐lingual expansion, P4 without ectostyle and with a well‐developed protocone, M1 bucco‐lingually elongated and double‐rooted, m1 with a larger talonid, and primitive mandible morphology. Thus, the population from Batallones‐1 constitutes a clearly different form from the genus Paramachaerodus, and we propose its inclusion in the genus Promegantereon  Kretzoi, 1938 , together with an upper canine from Crevillente‐2 (MN 11), very similar to those from Batallones‐1. In contrast, Pa. orientalis shows the following apomorphies: crenulated canines, P3 reduced in size and without disto‐lingual expansion, P4 with a clear ectostyle as well as a reduced, backwardly displaced protocone and with a rounded and single‐rooted M1. The species Pa. maximiliani is characterized by its slightly larger size, crenulated canines, very elongated P3 with a moderate disto‐lingual expansion and P4 and M1 similar to those of Pa. orientalis. Paramachaerodus orientalis is recorded at Puente Minero (MN 11, Teruel), Concud (MN 12, Teruel), Crevillente‐15, and Crevillente‐16 (both MN 12, Alicante), and Paramachaerodus maximiliani in Venta del Moro (MN 13, Valencia). The available data suggest that Pr. ogygia was present in the Iberian Late Vallesian and Early Turolian faunas (MN 10 and MN 11) but disappeared after that age. Paramachaerodus was present in the faunas throughout the Turolian, with the species Pa. orientalis and Pa. maximiliani, this latter being probably part of the same immigration event that occurred in the Late Turolian and involved other mammal taxa such as camelids and ursids.     

Brief communication: New primate remains from the Miocene of Namibia,Southern Africa

Miocene primates from southern Africa are extremely rare. For this reason we wish to place on record several interesting new fossil primate specimens recently recovered from the Miocene sites of Berg Aukas and Harasib in the Otavi Mountain region of northern Namibia. The new finds consist of a virtually complete atlas vertebra from Berg Aukas attributable to the hominoid Otavipithecus namibiensis and two teeth and four postcranial fragments from Harasib referrable to Cercopithecoidea. The atlas vertebra exhibits anatomical characteristics intermediate between those of modern cercopithecoids and hominoids which may be indicative of a transition from pronograde to orthograde postures. The cercopithecoid remains show that the earliest Old World monkeys known from southern Africa were small, approximately the size of vervet monkeys. These new specimens are important because they provide the first evidence relating to possible positional behaviors of Otavipithecus and the earliest fossil record of cercopithecoids from southern Africa. © 1996 Wiley-Liss, Inc.