Oligocene and Early Miocene gastropods from Kutch (NW India) document an early biogeographic switch from Western Tethys to Indo-Pacific

Shallow marine gastropod assemblages from Chattian, Aquitanian and Burdigalian sections in the Indian Kutch Basin are described. They provide insight into the composition and biogeographic relations of the gastropod assemblages at this junction between the Western Tethys and Proto-Indo-Pacific Ocean. For the first time, an improved biostratigraphy allows a clear separation of the assemblages, especially for the hitherto undifferentiated Early Miocene faunas. Throughout the Oligocene, about one-third of the species are also frequently found in the Western Tethys, documenting a passable Tethyan Seaway for nearshore molluscs. A considerable provincialism is evident as well. The expected turnover during the Early Miocene, due to the closing of the Tethyan Seaway, is reflected in the Miocene assemblages. Surprisingly, however, the cut appears very early, i.e. already during the Aquitanian, when the West–East interrelation drops to zero despite the passage having been open during this interval. In contrast, the Burdigalian assemblages witness a minor re-appearance of Western Tethys taxa, suggesting the re-establishment of rather ineffective migration pathways prior to the final closure of the Tethyan Seaway. Cerithium bermotiense and Lyria (Indolyria) maniyaraensis are introduced as new species.     

Dispersal of African mammals in Eurasia during the Cenozoic: Ways and whys

Several groups of mammals originated in Africa and then immigrated to Eurasia during some intervals of the Cenozoic, thus greatly contributing to the mammalian biodiversity in Eurasia. Nevertheless, the African components of Eurasian mammalian faunas have had variable success in their diversification and survival. The Afro-Arabian plate remained separated from Eurasia by the Tethyan seaway, which was definitely closed in the Burdigalian, some 20 myr ago. Before its closure, the marine barrier between the Afro-Arabian and Eurasian plates did not totally prevent mammalian exchanges between these landmasses, as documented by the arrival of rodents and primates in Africa in the late Paleocene-early Eocene, the dispersal of embrithopods on both sides of the Tethyan seaway during the Eocene, and the immigration of elephantoids from Africa to Asia in late Oligocene. These events seem to be restricted to some groups of mammals, which apparently had abilities to use sweepstake dispersal routes. The massive mammalian dispersal from Africa to Eurasia started sometimes in the early Miocene, involving several groups of African mammals, in particular proboscideans, hyracoids, tubulidentates, and anthropoids. This contribution discusses the timing of these events under the light of recent discoveries of Africa-originated mammals in Eurasia. The impact of the evolving paleogeography of the area situated between the Afro-Arabian and Eurasian plates on the mammalian dispersal is reconsidered. The dispersal of land mammals from Africa to Eurasia is controlled not only by the paleogeographic changes (sea level changes, dispersal routes, terrestrial bridges, etc.), but also by climatic factors that modified the environments of terrestrial mammals, favoring or not the occurrence of dispersal routes and/or the enlargement or restriction of climatic belts and biogeographic provinces to which these mammals were adapted. These questions are discussed taking into account the present knowledge of the record of the Africa-originated mammals in Eurasia during the Cenozoic times.     

Early Miocene reef- and mudflat-associated gastropods from Makran (SE-Iran)

A new gastropod fauna of Burdigalian (early Miocene) age is described from the Iranian part of Makran. The fauna comprises 19 species and represents three distinct assemblages from turbid water coral reef, shallow subtidal soft-bottom and mangrove-fringed mudflat environments in the northern Indian Ocean. Especially the reef-associated assemblage comprises largely new species. This is explained by the rare occurrence of reefs along the northern margin of the Miocene Indian Ocean and the low number of scientific studies dealing with the region. In terms of paleobiogeography, the fauna corresponds well to coeval faunas from the Pakistani Balochistan and Sindh provinces and the Indian Kathiawar, Kutch and Kerala provinces. During the early Miocene, these constituted a discrete biogeographic unit, the Western Indian Province, which documents the near complete biogeographic isolation from the Proto-Mediterranean Sea. Some mudflat taxa might represent examples of vicariance following the Tethys closure. The fauna also displays little connection with coeval faunas from Indonesia, documenting a strong provincialism within the Indo-West Pacific Region during early Miocene times. Neritopsis gedrosiana sp. nov., Calliostoma irerense sp. nov., Calliostoma mohtatae sp. nov. and Trivellona makranica sp. nov. are described as new species.     

Ecological implications of molluscan ontogenetic strategies - examples from aquatic ecosystems of the Cenozoic Iberian Peninsula

Molluscs from marginal marine and intra-continental basins of the Iberian Peninsula are described with special emphasis on the early ontogenetic shell formation, which reflects the embryogenesis and larval ecology. The fossils, covering a time span from the Early Oligocene to the Early Pleistocene, are compared to contemporaneous fossil faunas of the Mediterranean and Paratethys, and to extant Mediterranean faunas. Larval shells occur in bivalves and gastropods of Upper Tortonian coastal lagoons near Crevillente (Alicante), indicating marine larval stages and a connection of the adult habitat with the open sea. The euryhaline marginal marine gastropods display planktotrophic larval shells, which enable a marine distribution, but prevented generally euryhaline genera, such as Granulolabium and Terebralia (Cerithioidea, Potamididae), from colonizing continental saline ecosystems. The establishment of athalassic saline populations implies the preadaptational loss of planktotrophy. For the first time lecithotrophic larval development in an athalassic saline system is documented for Potamides gaudryi (Cerithioidea, Potamididae) from the late Middle to early Upper Miocene of the Duero Basin. With regard to the early ontogenetic development, P. gaudryi is distinguished from its possible descendant, the extant Potamides conicus (Blainville, 1829), which represents a direct developer that lacks any larval stage. In comparison to direct development, lecithotrophic larval development was advantageous in the colonization of temporary habitats, such as flood areas. Two different modes of direct development with hatching of crawling young are documented: Feeding on embryonic yolk until the hatching stage and adelphophagy. Adelphophagous embryonic development appears to be advantageous in neritids, thiarids and pulmonates that live in habitats with strong predation of juvenile fishes based on the advanced developmental stage and larger shell size of the hatchlings.     

Change in diversity,ecological significance and biogeographical relationships of the Mediterranean Miocene toothed whale fauna

The main evolutionary trend in the Mediterranean Miocene toothed whale fauna is related (1) to the change in diversity and (2) to the turnover in community structure. Diversity increases from Upper Aquitanian–Lower Burdigalian to Burdigalian–Langhian, when it reaches its maximum. Starting from this time, diversity decreases progressively. The Early Miocene (Upper Aquitanian–Lower Burdigalian) Mediterranean toothed whale fauna, as well as the extramediterranean ones, is characterised by a high number of endemic taxa and by the prevalence of longirostral forms living in estuarine-neritic environments. A more diversified fauna spreading in neritic and pelagic environments characterises the Burdigalian–Langhian age, while an increase in pelagic forms and the nearly complete disappearance of some archaic longirostral taxa is typical of the Serravallian–Messinian fauna. Decrease in diversity and disappearance of archaic longirostral taxa are also recorded, at more general scale, in the Late Miocene extramediterranean fossil bearing deposits. These events can be related to the progressive global climatic deterioration, starting from Middle Miocene. From a biogeographic point a view, we can outline some relationships between the Mediterranean and western North Atlantic Miocene faunas. Closer affinities are observed between the Baltringen fauna and the northern Atlantic one, because of the presence of the genera Pomatodelphis and Zarhachis (platanistids) in both areas. In the Miocene Mediterranean and in North Atlantic, the delphinids are apparently absent as well as other extant delphinoid groups even if erroneously recorded in the past.     

The spread of grass-dominated habitats in Turkey and surrounding areas during the Cenozoic: Phytolith evidence

The arrival of hipparionine horses in the eastern Mediterranean region around 11 Ma was traditionally thought to mark the simultaneous westward expansion of savanna vegetation across Eurasia. However, recent paleoecological reconstructions based on tooth wear, carbon isotopes, and functional morphology indicate that grasses played a minor role in Late Miocene ecosystems of the eastern Mediterranean, which were more likely dry woodlands or forests. The scarcity of grass macrofossils and pollen in Miocene floras of Europe and Asia Minor has been used to support this interpretation. Based on the combined evidence, it has therefore been suggested that Late Miocene ungulate faunal change in the eastern Mediterranean signals increased aridity and landscape openness, but not necessarily the development of grass-dominated habitats.

To shed new light on the Miocene evolution of eastern Mediterranean ecosystems, we used phytolith assemblages preserved in direct association with faunas as a proxy for paleovegetation structure (grassland vs. forest). We extracted phytoliths and other biogenic silica from sediment samples from well-known Early to Late Miocene ( 20–7 Ma) faunal localities in Greece, Turkey, and Iran. In addition, a Middle Eocene sample from Turkey yielded phytoliths and served as a baseline comparison for vegetation inference.

Phytolith analysis showed that the Middle Eocene assemblage consists of abundant grass phytoliths (grass silica short cells) interpreted as deriving from bambusoid grasses, as well as diverse forest indicator phytoliths from dicotyledonous angiosperms and palms, pointing to the presence of a woodland or forest with abundant bamboos. In contrast, the Miocene assemblages are dominated by diverse silica short cells typical of pooid open-habitat grasses. Forest indicator phytoliths are also present, but are rare in the Late Miocene (9–7 Ma) assemblages. Our analysis of the Miocene grass community composition is consistent with evidence from stable carbon isotopes from paleosols and ungulate tooth enamel, showing that C₄ grasses were rare in the Mediterranean throughout the Miocene. These data indicate that relatively open habitats had become common in Turkey and surrounding areas by at least the Early Miocene ( 20 Ma), > 7 million years before hipparionine horses reached Europe and arid conditions ensued, as judged by faunal data.     

A new dwarf boa (Serpentes, Booidea, 'Tropidophiidae') from the Early Oligocene of Belgium: a case of the isolation of Western European snake faunas

Falseryx neervelpensis sp. nov. (Booid-grade, 'Tropidophiidae') from the earliest Oligocene (MP 21) of Belgium is described on the basis of vertebrae coming from all major portions of the vertebral column. In its peculiar caudal osteology, the snake approaches the unique morphological pattern characteristic of the living Neotropical Tropidophiinae. This is the first time such a complete and informative vertebral column of a dwarf boa has been described. The genus Falseryx was absent from Western Europe in younger parts of the Oligocene, but reappeared at the end of the Early Miocene. This dispersal pattern provides additional evidence that in most phases of the Oligocene and Early Miocene Western European snake faunas were effectively isolated from possible influences from the East.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 152 , 393–406.     

Out of the Orient: Post‐Tethyan transoceanic and trans‐Arabian routes fostered the spread of Baorini skippers in the Afrotropics

The origin of taxa presenting a disjunct distribution between Africa and Asia has puzzled biogeographers for more than a century. This biogeographic pattern has been hypothesized to be the result of transoceanic long‐distance dispersal, Oligocene dispersal through forested corridors, Miocene dispersal through the Arabian Peninsula or passive dispersal on the rifting Indian plate. However, it has often been difficult to pinpoint the mechanisms at play. We investigate biotic exchange between the Afrotropics and the Oriental region during the Cenozoic, a period in which geological changes altered landmass connectivity. We use Baorini skippers (Lepidoptera, Hesperiidae) as a model, a widespread clade of butterflies in the Old World tropics with a disjunct distribution between the Afrotropics and the Oriental region. We use anchored phylogenomics to infer a robust evolutionary tree for Baorini skippers and estimate divergence times and ancestral ranges to test biogeographic hypotheses. Our phylogenomic tree recovers strongly supported relationships for Baorini skippers and clarifies the systematics of the tribe. Dating analyses suggest that these butterflies originated in the Oriental region, Greater Sunda Islands, and the Philippines in the early Miocene c. 23 Ma. Baorini skippers dispersed from the Oriental region towards Africa at least five times in the past 20 Ma. These butterflies colonized the Afrotropics primarily through trans‐Arabian geodispersal after the closure of the Tethyan seaway in the mid‐Miocene. Range expansion from the Oriental region towards the African continent probably occurred via the Gomphotherium land bridge through the Arabian Peninsula. Alternative scenarios invoking long‐distance dispersal and vicariance are not supported. The Miocene climate change and biome shift from forested areas to grasslands possibly facilitated geodispersal in this clade of butterflies.     

The tropical history and future of the Mediterranean biota and the West African enigma

Aim Since the opening of the Suez Canal in 1869, many tropical taxa from the Indo‐West Pacific (IWP) realm have entered the Mediterranean Sea, which is experiencing rising temperatures. My aims are: (1) to compare biogeographically this tropical transformation of the Mediterranean biota with the tropical faunas of the Mediterranean and adjacent southern European and West African seas during the Late Oligocene to Pliocene interval; (2) to infer the relative contributions of the tropical eastern Atlantic and IWP to the tropical component of the marine biota in southern Europe; and (3) to understand why West Africa is not now a major source of warm‐water species. Location Southern Europe, including the Mediterranean Sea, and the coast of tropical West Africa. Methods I surveyed the literature on fossil and living shell‐bearing molluscs to infer the sources and fates of tropical subgenus‐level taxa living in southern Europe and West Africa during the Late Oligocene to Pliocene interval. Results Ninety‐four taxa disappeared from the tropical eastern Atlantic (including the Mediterranean) but persisted elsewhere in the tropics, mainly in the IWP (81 taxa, 86%) and to a lesser extent in tropical America (36 taxa, 38%). Nine taxa inferred to have arrived in the tropical eastern Atlantic from the west after the Pliocene did not enter the Mediterranean. The modern West African fauna is today isolated from that of other parts of the marine tropics. Main conclusions Taxa now entering the Mediterranean through the Suez Canal are re‐establishing a link with the IWP that last existed 16 million years ago. This IWP element, which evolved under oligotrophic conditions and under a regime of intense anti‐predatory selection, will continue to expand in the increasingly warm and increasingly oligotrophic Mediterranean. The IWP source fauna contrasts with the tropical West African biota, which evolved under productive conditions and in a regime of less anti‐predatory specialization. Until now, the post‐Pliocene West African source area has been isolated from the Mediterranean by cold upwelling. If further warming should reduce this barrier, as occurred during the productive and warm Early Pliocene, the Mediterranean could become a meeting place for two tropical faunas of contrasting source conditions.     

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.     

Changes in the composition of the European snake fauna during the Early Miocene and at the Early / Middle Miocene transition

The members of the ‘modern’ snake families Colubridae, Viperidae and Elapidae (representatives of the family Colubridae appeared in Europe as the first) might have penetrated into the areas of Central Europe probably across the Mazury -Mazowsze continental bridge. The ‘modern’ families penetrated into West Europe across the Rhine Graben by several immigration waves. Based on the known marine incursions into the Rhine Graben and adjacent areas, a total of four distinct waves of dispersal of both the Asiatic and North American immigrants can be discerned within the Early Oligocene — Middle Miocene time span. The Early Miocene (MN 1–MN 2a) and the Early / Middle Miocene (MN 3–MN 4/5) dispersals of snakes are responsible for the final displacement of representatives of the ‘ancient’ family Boidae out of West and Central Europe. The Early and the Middle Miocene migrations have likewise initiated the evolution of the modern European snake fauna.     

Tethyan changes shaped aquatic diversification

The Tethys Ocean existed between the continents of Gondwana and Laurasia from the Triassic to the Pliocene. Analyses of multiple biogeographic and phylogenetic histories reveal that the subsequent breakup of the Tethys greatly influenced the distributions of many species. The ancestral Tethyan realm broke into five biogeographic provinces, including the present‐day East Pacific, West Atlantic, East Atlantic, Mediterranean Sea, and Indo‐West Pacific. Palaeogeographic maps illustrate the Mesozoic Atlantic opening, the Cenozoic closure of the Tethys, the Messinian Salinity Crisis, the mid‐Miocene closure of the Central American Seaway, and Quaternary geological changes. Further, we consider Cenozoic sea‐level changes and the formation of freshwater habitats. These reconstructions allow assessment of patterns of aquatic diversification for marine and freshwater animals, and comparison of vicariance and dispersal processes. Estimated divergence times indicate that fragmentation of the Tethys was responsible for the vicariant speciation of aquatic animals because these dates are consistent with associated tectonic events. The opening of the Atlantic Ocean during the Cretaceous is responsible for the earliest isolation between the West and East Atlantic. The mid‐Miocene closure of the Tethys, which blocked global equatorial currents, appears to have isolated the Atlantic/Mediterranean Sea and Indo‐West Pacific. Finally, formation of the Isthmus of Panama isolated East Pacific and West Atlantic marine organisms. Dispersals related to the Messinian Salinity Crisis and Quaternary sea‐level changes influenced population structuring. Tethyan changes affected marine habitats, created new freshwater habitats, inland caves and ancient lakes along the Alps and Himalayas, and influenced anchialine caves at the edge of the ancient sea. The extensive new habitats provided opportunities for colonisation and rapid diversification. Future work should focus on testing the biological impact of the series of Tethyan changes.     

步林在甘肃党河流域塔奔布鲁克地区的早期工作记录——经典脊椎动物化石地点与现代地层框架的解译

1931-1932年博格·步林在甘肃省西部塔奔布鲁克地区(又名党河地区,位于肃北县城西南)首次发现丰富的新生代哺乳动物化石。步林在塔奔布鲁克盆地中部燕丹图沟中找到三个小哺乳动物化石异常丰富的层位。三个层位紧靠在一起,其中产出的哺乳动物群被命名为燕丹图动物群。步林明确指出燕丹图动物群为晚渐新世。该动物群以后成为晚渐新世塔奔布鲁克哺乳动物期的典型动物群并沿用至今。     

The age of immigration of the vertebrate faunas found at Gargano (Apulia,Italy) and Scontrone (l’Aquila,Italy)

Recently a discussion is taking place about the Scontrone (l’Aquila) and Gargano (Apulia, Italy) mammal faunas and the age of their immigration. Mazza and Rustioni (2008) dated the Scontrone mammal fossils as Tortonian on the basis of their position in the Lithothamnium Limestone and came to the conclusion that some elements of the Scontrone and Gargano faunas must have colonised the area in Oligocene or Early Miocene times. Van den Hoek Ostende et al. (2009) disagreed with this interpretation and suggested a Late Miocene (10 Ma) age for the time of immigration. We think the arguments to place Scontrone in the Tortonian are not convincing. An analysis of the potential ancestors of each of the Gargano faunal components shows that a Messinian age for the immigration is fully compatible with the distribution of these ancestors in the European Miocene.     

Phylogenetic analysis informed by geological history supports multiple, sequential invasions of the Mediterranean Basin by the angiosperm family Araceae

Despite the remarkable species richness of the Mediterranean flora and its well-known geological history, few studies have investigated its temporal and spatial origins. Most importantly, the relative contribution of geological processes and long-distance dispersal to the composition of contemporary Mediterranean biotas remains largely unknown. We used phylogenetic analyses of sequences from six chloroplast DNA markers, Bayesian dating methods, and ancestral area reconstructions, in combination with paleogeographic, paleoclimatic, and ecological evidence, to elucidate the time frame and biogeographic events associated with the diversification of Araceae in the Mediterranean Basin. We focused on the origin of four species, Ambrosina bassii, Biarum dispar, Helicodiceros muscivorus, Arum pictum, subendemic or endemic to Corsica, Sardinia, and the Balearic Archipelago. The results support two main invasions of the Mediterranean Basin by the Araceae, one from an area connecting North America and Eurasia in the Late Cretaceous and one from the Anatolian microplate in western Asia during the Late Eocene, thus confirming the proposed heterogeneous origins of the Mediterranean flora. The subendemic Ambrosina bassii and Biarum dispar likely diverged sympatrically from their widespread Mediterranean sister clades in the Early-Middle Eocene and Early-Middle Miocene, respectively. Combined evidence corroborates a relictual origin for the endemic Helicodiceros muscivorus and Arum pictum, the former apparently representing the first documented case of vicariance driven by the initial splitting of the Hercynian belt in the Early Oligocene. A recurrent theme emerging from our analyses is that land connections and interruptions, caused by repeated cycles of marine transgressions-regressions between the Tethys and Paratethys, favored geodispersalist expansion of biotic ranges from western Asia into the western Mediterranean Basin and subsequent allopatric speciation at different points in time from the Late Eocene to the Late Oligocene.     

The beginnings of the Sargasso assemblage in the Tethys?

An assemblage of organisms, here termed “quasi-Sargasso”, occurs in the Oligocene menilite beds of the Carpathian flysch. It is suggested that this assemblage migrated into the Atlantic Ocean, probably during the Miocene. Its further evolution, which gave rise to the present-day Sargasso assemblage, took place in the middle Atlantic area.     

Neogene history of Sporolithon Heydrich (Corallinales,Rhodophyta) in the Mediterranean region

Most modern species of Sporolithon live in tropical and subtropical areas and only one species of the genus, S. ptychoides, occurs in the Mediterranean Sea. The scarce present-day populations of Sporolithon in the Mediterranean region are relics of a long history of the genus in this area since the Early Cretaceous. Analysis of data from the palaeontological literature, combined with the study of both fossil samples and Recent ones collected from Italy and Spain, shows that during the Miocene variations in the number of Sporolithon species in the Mediterranean region parallel changes in global temperature. After a maximum species richness in the Langhian (early Mid Miocene), coincident with the Miocene climatic optimum, the number of species decreased to just two before the Messinian Salinity Crisis. This marked decline follows the global cooling event that began at around 14 Ma. The closure of the connections of the Mediterranean region with the Indian Ocean during the Langhian left Mediterranean Sporolithon populations isolated from the main dispersal area of the genus. After the Messinian desiccation, a single species, S. ptychoides, re-invaded the Mediterranean Basin during the Early Pliocene and continues to inhabit this temperate sea today. The Atlantic Ocean is the most probable source of the re-invading Sporolithon plants.     

Systematics, biostratigraphy and evolutionary pattern of the Oligo-Miocene marine mammals from the Maltese Islands

An overview of the upper Oligocene-upper Miocene marine sediments outcropping in the Maltese Islands provides a detailed stratigraphical setting of several marine mammal assemblages. The studied fossil material collected within the entire sequence, is now kept in the National Museum of Natural History of Mdina (Malta). Nannoplankton analysis of some selected sections, where mammal remains have been discovered, is also undertaken. The fossil marine mammals, consisting mostly of isolated ear bones and teeth, are referred to cetaceans (both mysticetes and odontocetes), sirenians, and pinnipeds. The cetacean record evidences an evolutionary pattern that agrees with the Oligo-Miocene general trend, characterized by the progressive rarefaction and disappearance of archaic families (squalodontids, waipatiids, and, maybe, mammalodontids), and by the appearance and diversification of the extant families represented within younger strata (kogiids, pontoporiids and ziphiids). Pontoporiids, waipatiids, and tentatively mammalodontids are here reported for the first time in the Mediterranean, while the kogiid record represents the only sure Miocene evidence of this family in the Mediterranean. The geographical distribution of the mammalodontids and the waipatiids, based on the Maltese and extra-Mediterranean records, supports an open communication between the Proto-Mediterranean and the Indo-Pacific during the late Oligocene. Sirenians are represented by several dugongid pachyosteosclerotic rib fragments, collected from upper Oligocene through upper Miocene sediments. Pinnipeds are represented by a femur fragment from the Serravallian, referred to an indeterminate monachine, a phocid subfamily already reported from the Mio-Pliocene of the Mediterranean.     

Response of deep-sea benthic foraminifera to Miocene paleoclimatic events, DSDP Site 289

Changes in the Miocene deep-sea benthic foraminifera at DSDP Site 289 closely correlate to the climatically induced variations in deep and bottom waters in the Pacific Ocean. In early Miocene time, oxygen and carbon isotopes indicate that bottom waters were relatively warm and poorly oxygenated. Benthic foraminiferal assemblages are characterized by various species inherited from the Oligocene. Expansion of the Antarctic icecap in the early middle Miocene, 14–16 m.y. ago, increased oxygen isotope values, produced cold, more oxygenated bottom waters and lead to a turnover in the benthic foraminifera. An Oligocene—early Miocene assemblage was replaced by a cibicidoid-dominated assemblage. Some species became extinct and benthic faunas became more bathymetrically restricted with the increased stratification of deep waters in the ocean. In mid-Miocene time, Epistominella exigua and E. umbonifera, indicative of young, oxygenated bottom waters, are relatively common at DSDP Site 289. Further glacial expansion 5–9 m.y. ago lowered sealevel, increased oceanic upwelling and associated biological productivity and intensified the oxygen minima. Abundant hispid and costate uvigerines become a dominant faunal element at shallow depths above 2500 m as E. umbonifera becomes common to abundant below 2500 m. By late Miocene time, benthic faunas similar in species composition and proportion to modern faunas on the Ontong-Java plateau, had become established.