Systematic revision of Siwalik Rhizomyidae (Rodentia)

Larges samples of fossil rhizomyids from the Siwalikmolasse deposits of the Potwar Plateau, Pakistan, enable reevaluation of fossil taxa and their evolutionary relationships. Siwalik Rhizomyidae constitute a basis for analysis of poorly represented rhizomyids from other areas in Asia and contribute to biochronological correlation. The extant Tachyoryctinae and Rhizomyinae diverged during the early history of the family and diagnoses of the subfamilies are amended to include extinct genera. Siwalik tachyoryctines include Kanisamys, Protachyoryctes, Rhizomyides, and Eicooryctes nov. gen.; rhizomyines are Brachyrhizomys and Anepsirhizomys nov. gen. Kanisamys, the earliest Siwalik rhizomyid, appeared by about 13 Ma and evolved slowly, displaying stasis in K. sivalensis. Brachyrhizomys evolved from Kanisamys by 9.5 Ma and radiated rapidly. B. nagrii displays rapid increase in size through time. About 7 Ma, Kanisamys and Brachyrhizomys became extinct locally and Protachyoryctes, Eicooryctes, and Rhizomyides appeared, perhaps in response to increasing aridity. After 5.5 Ma, rhizomyids became uncommon in the Potwar and Anepsirhizomys (3.0 Ma) constitutes the latest record of Rhizomyidae in Pakistan.     

Tragoportax cf. rugosifrons (Schlosser, 1904) from the late Miocene of Cessaniti (Southern Italy)

This paper describes remains attributable to Tragoportax cf. rugosifrons (Schlosser, 1904) found in the late Miocene site of Cessaniti (Vibo Valentia, Calabria) and the surrounding area. The studied specimens come from the Clypeaster sandstones, included in a marine/fluvial succession dated between 8 and 7.2 Ma. At Cessaniti, Tragoportax is associated with Stegotetrabelodon syrticus Petrocchi, 1941; Samotherium cf. boissieri Forsyth-Major, 1888; Bohlinia cf. attica Matthew, 1929; and an undetermined Rhinocerotid still under study. The genus Tragoportax was common in Eurasia and Africa during the late Miocene. The occurrence of Tragoportax cf. rugosifrons at Cessaniti confirms the peculiarity of the assemblage, with its association of species of North African and Pikermian (Greco-Iranian bioprovince) affinities.     

Pliocene Scelidotheriinae (Xenarthra,Tardigrada) from the Pampean region of Argentina: Morphology,chronology, and comments on the diversity of the subfamily

Scelidotheriinae (Mammalia, Pilosa) is a xenarthran mylodontid clade recorded in much of South America from the middle Miocene to the early Holocene. However, studies of their Neogene representatives are still scarce compared to their Quaternary representatives. The main goal of this contribution is to report new remains of Proscelidodon rothi (Ameghino, 1908) found in the basal levels of the El Polvorín Formation (Pliocene), cropping out near the city of Olavarría (Buenos Aires Province, Argentina). On the basis of the faunistic association, these deposits were assigned to the late Pliocene. The new materials include a skull, a mandible, and a calcaneal fragment (Xen-121), which enhance the characterization of this taxon, and allow more precise comparisons with other Neogene and Quaternary Scelidotheriinae. This new record extends the biochron and geographic distribution of the species, until now recorded in the latest Pliocene (Marplatan Stage/Age, Vorohuean Subage) of Olavarría.     

Geopalaeontological setting,chronology and palaeoenvironmental evolution of the Baccinello-Cinigiano Basin continental successions (Late Miocene,Italy)

The Latest Miocene succession of the Baccinello-Cinigiano Basin in southern Tuscany (Italy) recorded a faunal turnover documenting the extinction of an older, insular, endemic faunal complex characterised by the extinct ape Oreopithecus bambolii and the setting of a new, continental, European faunal complex including the colobine monkey Mesopithecus. A similar turnover pattern (Late Miocene ape/Latest Miocene Cercopithecidae) is generally observed in Late Miocene continental successions of Eurasia, from Spain to central Europe, Southwest Europe, the near East, and Southwest Asia. Abundant literature reports that the Late Miocene Eurasian hominoid primate distribution closely tracks the climatic/environmental changes occurring during the 12–9 Ma interval, until their extinction in western Europe. In the primate record, the dispersion of Cercopithecidae and the contraction of hominids is interpreted as an event depicting a pattern of “continentalisation” in the Old World. The sedimentary succession of the Baccinello-Cinigiano basin, one of the longest continuous vertebrate-bearing continental successions in the Neogene Italian record, contributes to the debate on this hypothesis. This paper provides an overview of the main characteristics of the sedimentary succession, the chronological constraints (biochronology, radiometric datings, magnetostratigraphy), and the palaeoenvironmental evolution as derived from palaeobiological approaches and from the study of stable carbon and oxygen isotope contents along the entire sedimentary succession. The 2 myr geological history of the Baccinello Cinigiano Basin, which documents the evolutionary history of Oreopithecus and associated faunas, does not have a direct relation with the event of the Messinian Salinity Crisis. The evolutionary history of Baccinello-Cinigiano Basin and its palaeontological record have been mainly driven by the regional tectonism and palaeogeographic changes that affected the northern Tyrrhenian regions in Late Miocene (Latest Tortonian–Messinian) times.     

The first representative Pliocene assemblages of passerine birds in Asia (Northern Mongolia and Russian Transbaikalia)

Passerine birds from two late Pliocene localities, Shaamar in northern Mongolia and Beregovaya in Transbaikalia (East Russia), represent the first known Neogene fossil assemblages of perching birds from Asia. A total of 11 families and 15 taxa are recognized, including a new genus and species of bunting, Pliocalcarius orkhonensis, apparently related to the Longspurs, and new species within the genera Hirundo and Rhodospiza, which are described. The avifauna is composed of open land and bush dwellers; it contains a few arid species, representing the oldest finds of the corresponding genera or lineages. This may indicate that the living Asian arid avian complex originated in Central Asia in the Neogene. Several genera (Rhodospiza, Paradoxornis) are documented in this paper for the first time in pre-Pleistocene deposits. Report of Calcarius is the first pre-Holocene evidence of the occurrence of this genus in Asia, which together with the remains of Synaptomys lemmings at Shaamar, illustrates faunal interchanges between North America and Asia in the Pliocene.     

Simimys and origin of the cricetidae (Rodentia: Muroidea)

Simimys is a late Eocene and earliest Oligocene genus that shares dipodoid (zygoma) and muroid (dental) characters. The Oligocene record of dipodoid rodents includes Plesiosminthus from middle Oligocene deposits of Asia and late Oligocene deposits of Europe. The Oligocene record of muroid rodents includes at least two genera (Eucricetodon and (Cricetops)) from Asia, six genera (Eucricetodon, Pseudocricetodon, Melissiodon, Paracricetodon, Heterocricetodon and Adelomyorion)) from Europe, and three genera (Eumys, Scottimus and Nonomys)) from North America. The known record, as given above, suggests that Siminys is the earliest and most primitive genus with muroid affinities; it also implies that muroid rodents were derived from unknown Eocene dipodoid rodents.The Oligocene cricetid rodents display progressive expansion and inclination of the anterior plate of the zygoma. These changes in the zygoma probably reflect evolutionary stages in the development of a myomorphous zygoma from an hystricomorphous zygoma. Changes in the zygoma apparently took place at different rates and times in Asia. Europe, and North America; they probably reflect differenciation of Cricetodontinae in Palaearctica, and Eumyinae in Nearctica during the Oligocene.     

Phytogeographical Implication of Bridelia Will. (Phyllanthaceae) Fossil Leaf from the Late Oligocene of India

Background

The family Phyllanthaceae has a predominantly pantropical distribution. Of its several genera, Bridelia Willd. is of a special interest because it has disjunct equally distributed species in Africa and tropical Asia i.e. 18–20 species in Africa-Madagascar (all endemic) and 18 species in tropical Asia (some shared with Australia). On the basis of molecular phylogenetic study on Bridelia, it has been suggested that the genus evolved in Southeast Asia around 33±5 Ma, while speciation and migration to other parts of the world occurred at 10±2 Ma. Fossil records of Bridelia are equally important to support the molecular phylogenetic studies and plate tectonic models.

Results

We describe a new fossil leaf of Bridelia from the late Oligocene (Chattian, 28.4–23 Ma) sediments of Assam, India. The detailed venation pattern of the fossil suggests its affinities with the extant B. ovata, B. retusa and B. stipularis. Based on the present fossil evidence and the known fossil records of Bridelia from the Tertiary sediments of Nepal and India, we infer that the genus evolved in India during the late Oligocene (Chattian, 28.4–23 Ma) and speciation occurred during the Miocene. The stem lineage of the genus migrated to Africa via “Iranian route” and again speciosed in Africa-Madagascar during the late Neogene resulting in the emergence of African endemic clades. Similarly, the genus also migrated to Southeast Asia via Myanmar after the complete suturing of Indian and Eurasian plates. The emergence and speciation of the genus in Asia and Africa is the result of climate change during the Cenozoic.

Conclusions

On the basis of present and known fossil records of Bridelia, we have concluded that the genus evolved during the late Oligocene in northeast India. During the Neogene, the genus diversified and migrated to Southeast Asia via Myanmar and Africa via “Iranian Route”.     

Famennian (Late Devonian) ammonoids from the Ouarourout section (Saoura Valley,Algeria)

In North Africa, the ammonoids constitute an important part of the Devonian marine macro-invertebrates. New material has been recently collected in the ‘Argiles de Marhouma’ Formation from several sections located near Béni-Abbès in the Saoura Valley (Ougarta Mountains, Algerian Sahara). Red nodular limestones (i.e., ‘griotte’ facies) characterize this formation rich in ammonoids. The assemblages from the Ouarourout section are composed of Goniatitida and Clymeniida constituting 30 species and 19 genera belonging to 11 families, with Cyrtoclymenia, Erfoudites, Kosmoclymenia, Planitornoceras, Platyclymenia, and Prionoceras, as the most abundant genera. The taxonomic comparison and the biostratigraphic correlations are made by considering the assemblages recorded in the Anti-Atlas of Morocco, which are very close to those observed in Algeria. In the Ouarourout area, the ‘Argiles de Marhouma’ Formation contains middle and late Famennian ammonoid assemblages and three biostratigraphic intervals can be identified: Planitornoceras, Platyclymenia and Medioclymenia.     

The Biogeographic South-North Divide of Polygonatum (Asparagaceae Tribe Polygonateae) within Eastern Asia and Its Recent Dispersals in the Northern Hemisphere

Eastern Asia (EA) is a key region for the diversification of flowering plants in the Northern Hemisphere, but few studies have focused on the biogeographic history within EA in the context of the other northern continents. Polygonatum is an important medicinal genus widely distributed in the Northern Hemisphere with its highest species richness in EA, and it represents an excellent model for studying the evolution of biogeographic patterns in this region. Divergence time estimation was used to examine the biogeographic history of Polygonatum based on nuclear ITS and four plastid sequences (rbcL, matK, psbA–trnH and trnC–petN) from 30 Polygonatum species and 35 outgroup taxa. The ancestral area of Polygonatum and subsequent dispersal routes were inferred using Bayes-Lagrange. Polygonatum was estimated to have originated in southern EA during the middle Miocene (14.34–13.57 Ma) with subsequent south-to-north expansion in the late Miocene. Multiple intercontinental dispersal events were inferred between EA and Europe or North America, and all of them have occurred recently in the late Miocene to Pliocene. The separation of Polygonatum into the south and north lineages and their subsequent diversifications in the late Miocene supports the existence of a biogeographic divide between the northern and southern parts of EA that also coincides with the retreat and redevelopment of the arid zone in EA in the Neogene. Our results demonstrate the complexity of biogeographic history of Polygonatum in the Northern Hemisphere including early vicariance followed by frequent and recent dispersals in the Neogene.     

The Anatidae and Scolopacidae (Aves: Anseriformes,Charadriiformes) from the late Neogene of Gargano,Italy

The late Neogene (MN13-14) fissure fillings found in the limestone quarries near Apricena (Foggia, Southern Italy) contain a well-diversified fossil bird assemblage. Most of the bird taxa show endemic characteristics following the high degree of insularity of the whole vertebrate assemblage. In addition to the endemic taxa, some non-endemic forms are present, mostly only recently found after the still ongoing revision of the whole bird remains. Here the remains of Anatidae and Scolopacidae are presented. This analysis reveals the occurrence of at least two taxa of Anatidae, Anas velox and Anatidae indet., and two taxa of Scolopacidae, Calidris sp. and an Scolopacidae indet. In addition, some remains are determined as Charadriiformes indet., but they probably represent more than one taxon, even if their bad status of preservation does not allow any further consideration. The detailed study of these remains and their comparison with the other European Neogene taxa already described is carried out.     

A revision of fossil Rhizomyid rodents from Northern India and their correlation to a Rhizomyid biochronology of Pakistan

Based on the biochronology of rhizomyid rodents of Northern Pakistan, fossil horizons in the Siwaliks of Northern India can be correlated to the late Neogene sequence of Pakistan with greater precision than previously possible. The indian Siwaliks do not include records of the most primitive Rhizomyidae, but rhizomyid faunas of India display strong affinity with those of Pakistan in the late Miocene. Pliocene differentiation of rhizomyid faunas in India and Pakistan may reflect ecological partitioning within the Indian subcontinent. Use of the term «Nagri for deposits at Haritalyangar suggests an erroneous equivalence in age to the Nagri stratotype in Pakistan.     

Phytosaur remains (Reptilia,Thecodontia)from the Upper Triassic of North-Eastern Thailand

The recently discovered late Triassic vertebratelocality at Chulabhorn Dam (North-Eastern Thailand) has yielded incomplete remains (jaw fragments and teeth) of phytosaurs, which are apparently indicative of a form related to Belodon and Rutiodon. They can be interpreted as showing that in the late Triassic North-Eastern Thailand was already biogeographically part of Laurasia.     

The taxonomy,chronostratigraphy and paleobiogeography of glyptosaurine lizards (Glyptosaurinae,Anguidae)

Glyptosaurine lizards (Glyptosaurinae, Anguidae) are an extinct group of heavily armored lizards known from North America, Europe and Asia. Glyptosaurine lizards, taxa that possess fully developed tuberculated dermal armor, appear to have been established in North America by late early Puercan time (To3). “Proxestops,” a taxon distinguished by a combination of vermiculate and tuberculated osteoderm sculpturing, is considered to be a non-glyptosaurine, a sister taxon of the Glyptosaurinae. Known from only fragmentary remains, its wide chronostratigraphic distribution suggests that “Proxestops” is a form genus that, in all probability, represents more than one taxon, that ranges from the middle Paleocene to the early Eocene of North America. Moreover, the taxa Odaxosaurus piger, Parodaxosaurus sanjuanensis and “Proxestops” are best considered “proto-glyptosaurines”. “Melanosaurins” and glyptosaurins were well-established by the early Eocene, especially in North America, and are here documented by their type species and chronostratigraphic levels. Both tribes are present in Europe (MP7), too, but the record is not as estensive as that of North America. The North American taxon Gaultia silvaticus (Wa0) is transitional between a “melanosaurin” and glyptosaurin. Because it lacks the well-defined hexagonal osteoderms that characterize the Glyptosaurini, it is removed from that group and considered to be a “melanosaurin”. The “melanosaurin” taxon “Xestops” savagei (Wa4–Wa6) cannot be referred to Xestops (Br2) based on non-corresponding elements and because superficial similarity does not justify assignment to this taxon. Arpadosaurus sepulchralis (Wa6?), whose holotype is a fragmentary right frontal, is considered a subjective junior synonym of A. gazinorum, based on minor differences in the epidermal scale pattern that probably represent individual variation. “Glyptosaurus” agmodon (Wa6?), based on a partial right maxilla, cannot be referred to Glyptosaurus (sensu stricto), and the material upon which this taxon is based bears strong resemblance to material identified as cf. “?Paraglyptosaurus” yatkolai (Wa5–Wa6). “Glyptosaurus” rhodinos (Wa5) is based on an incomplete parietal, and its reference to Glyptosaurus is considered problematic. Eoglyptosaurus donohoei (Wa7) is probably valid and is re-established here. Glyptosaurus (sensu stricto) is known solely from the middle Eocene (Br2) by G. sylvestris. Dimetoposaurus wyomingensis (Br3) is removed from Xestops vagans because its synonymy was based on superficial similarities. Helodermoides tuberculatus, the largest and last glyptosaurin (Ch3), is restricted to the Chadronian of North America. Only the “melanosaurin” Peltosaurus granulosus (Or2–Or3), which includes the species P. abbotti, seems to have crossed the Eocene-Oligocene boundary, and appears to be largely restricted to the Orellan, but extended into the Arikareean. European glyptosaurines are also represented by both glyptosaurins and “melanosaurins” early in the Eocene (MP7). Placosauriops-like “melanosaurins” are known from Dormaal (MP7), and the glyptosaurin taxon?Placosaurus ragei occurs at the same level. “Placosauriops abderhaldeni” has been identified from the Grube Messel (MP11), but this assignment remains dubious because the species has not been adequately diagnosed, and the holotype species is from the Geiseltal (MP13), which is some 4.5 million years younger. Placosauriops weigelti (MP13) is the only valid species of this genus. Paraxestops stehlini (MP14) is not referable to the North American taxon Xestops, and its relationship to Placosauriops has not been studied. The late Eocene glyptosaurins Placosaurus estesi (MP17) and P. rugosus (MP18) are the last glyptosaurines known from Europe and appear to have gone extinct at the Eocene-Oligocene boundary, casulties perhaps of the “Grande Coupure”. Asian glyptosaurines are known solely from one species, Stenoplacosaurus mongoliensis, from the middle Eocene (Sharamurunian) of China. Glyptosaurines most likely originated in North America, diversified by late Paleocene time, and rapidly spread across the North Atlantic into Europe by the early Eocene. Both “melanosaurins” and glyptosaurins took a foothold in Europe by the early Neustrian, but the glyptosaurins, aside from one occurrence (Dormaal, MP7), were conspicuously absent for most of Neustrian through early Robiacian time. In North America, glyptosaurins diversified during the early and middle Eocene, while in Europe small “melanosaurins” were a prominent part of the paleoherpetofauna, and glyptosaurins are unknown for most of the Neustrian through the Geiseltalian, in both the fossilferous Lagerstätten of Messel and Geiseltal. Stenoplacosaurus is the only known glyptosaurin glyptosaurine from Asia, and its abrupt appearance during the late Eocene suggests the possiblity of a Beringian dispersal from North America into Asia.     

Evolution of bird communities in the Neogene of Central Asia,with a review of the Neogene fossil record of Asian birds

A complete taxonomic review of Neogene birds of continental Asia is provided. To date, avifauna from the latter half of the Miocene and Pliocene of Central Asia (Mongolia and adjacent regions of Inner Asia) are most thoroughly investigated. Available data enable a reconstruction of successive replacement of Early and Middle Miocene avifaunas by communities of the Recent type. Middle Miocene avifaunas of Mongolia include a great number of extinct genera and species, many of which were widespread in Eurasia. Extant genera became dominant in the Late Miocene and taxa close to living species appear in the Late Pliocene fossil record. Late Pliocene communities of birds of Central Asia were complex in genesis, composed of Miocene relicts (Struthio), immigrants from the European regions of the Palearctic (phasianid Plioperdix), North American immigrants (Calcarius), and also autochthonous elements, the origin of which is apparently connected with the arid belt of Central Asia (diverse passerines).     

Amphibians and squamate reptiles from the late Pleistocene of the “Caverne Marie-Jeanne” (Hastière-Lavaux,Namur, Belgium): Systematics,paleobiogeography, and paleoclimatic and paleoenvironmental reconstructions

Archeological sites usually provide important information about the past distribution of small vertebrate fauna, and by extension about past terrestrial environments and climate in which human activities took place. In this context, Belgium has an interesting location in northwestern Europe between the fully studied zooarcheological records of Germany and England. We present here the revision of the late Pleistocene (Marine Isotope Stages 3 and 2) collection of the “Caverne Marie-Jeanne” (Hastière-Lavaux, Namur), studied by Jean-Claude Rage in the 1970s and the revision of the whole “indeterminate” small vertebrate materials from the “Caverne Marie-Jeanne” stored in the Royal Belgian Institute of Natural Sciences (RBINS) Quaternary collections in search of more herpetofaunal remains. It is now by far the largest late Pleistocene collection at RBINS with more than 20,500 recognized bones of amphibians and reptiles and covering the last 60,000 years. The faunal list comprises two urodeles (Lissotriton gr. L. vulgaris and Salamandra salamandra), four anurans (Bufo gr. B. bufo-spinosus, Epidalea calamita, Rana temporaria and Rana cf. R. arvalis), three lizards (Lacerta cf. L. agilis, Zootoca vivipara and Anguis gr. A. fragilis), and three snakes (Natrix gr. N. natrix, Coronella austriaca, and Vipera berus). This study represents the first fossil record in Belgium for L. gr. L. vulgaris, R. arvalis, Z. vivipara, N. gr. N. natrix and C. austriaca. As a whole, this assemblage suggests a patchy humid landscape under colder and dryer climatic conditions in comparison with present ones. This study also underlines the necessity of a primary separation in larger taxonomical categories by the specialist itself.     

Presence of Hypolagus Dice, 1917 (Mammalia, Lagomorpha) in the Neogene of the Balearic Islands (Western Mediterranean): Description of Hypolagus balearicus nov. sp.

In this paper we describe a new species of Hypolagus coming from an early Pliocene karstic deposit near Caló den Rafelino (Manacor, Mallorca). It represents the westernmost European record for the genus. The faunal assemblage of this deposit represents an early phase of the second insular faunal episode of Mallorca, related with the Messinian regressive episode (latest Miocene). Although the postcranial are relatively robust, the size of all these elements is included in the range of continental leporids. It is unknown if they display significant allometric changes, as occurred in other insular leporids. p3 is small and displays a markedly trapezoidal outline and a deep hypoflexid (between 54 and 56% of the total tooth width). The presence of Hypolagus in the Neogene of the Balearic Islands is in agreement with the faunal scenario for the European continent during the late Miocene and the Pliocene, where this genus is abundant and widely distributed.     

Dragon's Paradise Lost: Palaeobiogeography,Evolution and Extinction of the Largest-Ever Terrestrial Lizards (Varanidae)

Background

The largest living lizard species, Varanus komodoensis Ouwens 1912, is vulnerable to extinction, being restricted to a few isolated islands in eastern Indonesia, between Java and Australia, where it is the dominant terrestrial carnivore. Understanding how large-bodied varanids responded to past environmental change underpins long-term management of V. komodoensis populations.

Methodology/Principal Findings

We reconstruct the palaeobiogeography of Neogene giant varanids and identify a new (unnamed) species from the island of Timor. Our data reject the long-held perception that V. komodoensis became a giant because of insular evolution or as a specialist hunter of pygmy Stegodon. Phyletic giantism, coupled with a westward dispersal from mainland Australia, provides the most parsimonious explanation for the palaeodistribution of V. komodoensis and the newly identified species of giant varanid from Timor. Pliocene giant varanid fossils from Australia are morphologically referable to V. komodoensis suggesting an ultimate origin for V. komodoensis on mainland Australia (>3.8 million years ago). Varanus komodoensis body size has remained stable over the last 900,000 years (ka) on Flores, a time marked by major faunal turnovers, extinction of the island''s megafauna, the arrival of early hominids by 880 ka, co-existence with Homo floresiensis, and the arrival of modern humans by 10 ka. Within the last 2000 years their populations have contracted severely.

Conclusions/Significance

Giant varanids were once a ubiquitous part of Subcontinental Eurasian and Australasian faunas during the Neogene. Extinction played a pivotal role in the reduction of their ranges and diversity throughout the late Quaternary, leaving only V. komodoensis as an isolated long-term survivor. The events over the last two millennia now threaten its future survival.     

Paracamelus (Mammalia,Camelidae) remains from the late Messinian of Italy: insights into the last camels of western Europe

In this paper we describe fossil remains of an indeterminate species of the genus Paracamelus (Artiodactyla, Camelidae) from the Messinian post-evaporitic deposits (5.55–5.40 Ma) of Verduno (Piedmont, NW Italy). Camelins dispersed into Eurasia from North America in the late Miocene and almost instantaneously spread in western Europe and Africa. The size and morphology of the fossils found at Verduno are consistent in with those of Paracamelus, the earliest Old World camelin. Up to now, the only fossil camels recovered in the Neogene of Western Europe have been found at Venta del Moro and Librilla in Spain at 6.2 Ma. The remains from Verduno represent the first evidence of a camelin in the Neogene of Italy and they considerably expand the paleobiogeographic range of the Old World early camelins. The presence of a camelid at Verduno reinforces and confirms the importance of the fossiliferous deposits of NW Italy in defining the complex paleobiogeographic patterns of Europe during the Messinian, at the end of the Messinian salinity crisis.