Re-evaluation of the morphology and phylogeny of Diplocynodon levantinicum Huene & Nikoloff, 1963 and the stratigraphic age of the West Maritsa coal field (Upper Thrace Basin, Bulgaria).

Diplocynodon levantinicum Huene & Nikoloff, 1963 was described based on few bone fragments from the West-Maritsa lignite basin of Central Bulgaria. Huene & Nikoloff, 1963 assumed a late Pliocene age, implying that this species represents the stratigraphically youngest crocodilian of Europe. In this current study, we re-evaluate the stratigraphy of the West-Maritsa Basin and conclude a late Oligocene age of ~26 Ma for the Kipra coal-seam, the fossiliferous horizon. Furthermore, topotypical and undescribed D. levantinicum specimens are accessible now and allowed for a deeper taxonomic and phylogenetic analysis. A comparison with other Diplocynodon species reveals D. levantinicum as a valid species, having (1) a long suborbital fenestra, (2) a very short dentary symphysis, (3) a large gap between the first and second dentary alveolus, (4) an occlusion pit in line with the tooth row posterior to the 14th dentary alveolus, (5) a sulcus lateral to the glenoid fossa and, (6) a lingual foramen for the articular artery situated entirely on the surangular. The phylogenetic analyses find D. levantinicum deeply nested inside the Diplocynodontinae subfamily. After the disappearance of the Paratethyan influence (Solenovian regional stage) in the Upper Thrace Basin this species has roamed during the late Oligocene extensive freshwater lake and swamp ecosystems represented by the Maritsa Formation. Diplocynodon levantinicum represents the only nominal Diplocynodon taxon of late Oligocene (Chattian) age.

The exceptionally high diversity of small carnivorans from the Late Miocene hominid locality of Hammerschmiede (Bavaria, Germany).

The present study deals with new material of carnivorans (Mustelidae, Mephitidae, Ailuridae, Potamotheriinae and Viverridae) from the basal Tortonian (Late Miocene, late Astaracian) hominid-bearing locality of Hammerschmiede (Bavaria, Germany). The small carnivoran fauna includes 20 species belonging to nine different subfamilies (Guloninae, Lutrinae, Mellivorinae, Potamotheriinae, Leptarctinae, Mephitinae, Simocyoninae, Genettinae and Viverrinae). The identified forms include: "Martes" sansaniensis, "Martes" cf. munki, "Martes" sp., Circamustela hartmanni n. sp., Laphyctis mustelinus, Guloninae indet., Eomellivora moralesi, Vishnuonyx neptuni, Paralutra jaegeri, Lartetictis cf. dubia, Trocharion albanense, Palaeomeles pachecoi, Proputorius sansaniensis, Proputorius pusillus, Alopecocyon goeriachensis, Simocyoninae indet., Potamotherium sp., Semigenetta sansaniensis, Semigenetta grandis and Viverrictis modica. The new species Circamustela hartmanni n. sp. is differentiated from the other members of the genus by its small size and the morphology of its dental cusps in the upper and lower carnassials. This is one of the highest reported taxonomic diversities for fossil small carnivorans in the Miocene of Europe, including also first and last occurrences for several genera and species. Additionally, the assemblage comprises some rare taxa such as Palaeomeles pachecoi and Eomellivora moralesi. An ecomorphological comparison of the discovered taxa reveals possible cases of competition and niche partitioning.

Unexpected cranial sexual dimorphism in the tragulid Dorcatherium naui based on material from the middle to late Miocene localities of Eppelsheim and Hammerschmiede (Germany).

Tragulids, chevrotains or mouse deer, were common faunal elements during the Miocene. During that time, Dorcatherium was the most abundant genus, with D. naui being the first described species. Besides their abundance, until recently only very limited cranial material was available for investigation. Here we present a redescription of the first complete skull of D. naui from the middle to late Miocene locality of Eppelsheim, Germany, based on micro-computed tomography. Furthermore, we present a description and comparison of two additional, new skulls of D. naui from the late Miocene hominid locality Hammerschmiede, Germany. Within Dorcatherium, so far, only three other complete skulls are known, all belonging to D. crassum. A comparison between the three skulls of D. naui and the already known skulls of D. crassum shows that these two species differ in morphological features of the skull, such as laterally facing orbitae, separation of supraorbital foramen from supraorbital groove by a bony bridge, well-developed parietal plateau, prominent nuchal tubercle, less-developed nuchal crests, and the presence of an occipital crest. Moreover, two different osteological morphotypes are present in the skulls of D. naui that can be interpreted as a previously unknown sexual dimorphism. Very similar features are observed in D. crassum, which can be likewise related to the same dimorphism. However, males of D. naui differ from males of D. crassum by the presence of frontoparietal bulges, which were probably used for sexual display and during male-male combats in males of D. naui. For the first time, sexual dimorphism in Dorcatherium is described based on skull characteristics, which are, so far, unknown from any other fossil or extant tragulid.

Age constraints for the Trachilos footprints from Crete.

We present an updated time frame for the 30 m thick late Miocene sedimentary Trachilos section from the island of Crete that contains the potentially oldest hominin footprints. The section is characterized by normal magnetic polarity. New and published foraminifera biostratigraphy results suggest an age of the section within the Mediterranean biozone MMi13d, younger than ~ 6.4 Ma. Calcareous nannoplankton data from sediments exposed near Trachilos and belonging to the same sub-basin indicate deposition during calcareous nannofossil biozone CN9bB, between 6.023 and 6.727 Ma. By integrating the magneto- and biostratigraphic data we correlate the Trachilos section with normal polarity Chron C3An.1n, between 6.272 and 6.023 Ma. Using cyclostratigraphic data based on magnetic susceptibility, we constrain the Trachilos footprints age at ~ 6.05 Ma, roughly 0.35 Ma older than previously thought. Some uncertainty remains related to an inaccessible interval of ~ 8 m section and the possibility that the normal polarity might represent the slightly older Chron C3An.2n. Sediment accumulation rate and biostratigraphic arguments, however, stand against these points and favor a deposition during Chron C3An.1n.

The large-sized darter Anhinga pannonica (Aves, Anhingidae) from the late Miocene hominid Hammerschmiede locality in Southern Germany.

We report fossils of the darter Anhinga pannonica Lambrecht, 1916 from two late Miocene (Tortonian, 11.62 and 11.44 Ma) avifaunas in Southern Germany. The material from the hominid locality Hammerschmiede near Pforzen represents the most comprehensive record of this species and includes most major postcranial elements except for the tarsometatarsus. We furthermore show that the putative cormorant Phalacrocorax brunhuberi (von Ammon, 1918) from the middle Miocene of Regensburg-Dechbetten is another, previously misclassified, record of A. pannonica, and this may also be true for early Miocene fossils described as P. intermedius Milne-Edwards, 1867. A. pannonica was distinctly larger than extant darters and reached the size of A. grandis from the late Miocene of North America. We detail that only fossils from the Miocene of Europe and Africa can be referred to A. pannonica, whereas putative records from Asia fall within the size range of extant darters. A. pannonica appears to have been a long-living species (16 to 6 Ma) with an extensive distribution from the equator to the northern mid-latitudes. The extinction of large-sized darters in Europe is likely to have been due to climatic cooling in the late Neogene, but the reasons for their disappearance in Africa and South America remain elusive.

A new alligatoroid from the Eocene of Vietnam highlights an extinct Asian clade independent from extant Alligator sinensis.

During systematic paleontological surveys in the Na Duong Basin in North Vietnam between 2009 and 2012, well-preserved fossilized cranial and postcranial remains belonging to at least 29 individuals of a middle to late Eocene (late Bartonian to Priabonian age (39-35 Ma)) alligatoroid were collected. Comparative anatomical study of the material warrants the diagnosis of a new taxon, Orientalosuchus naduongensis gen. et sp. nov. The combined presence of an enlarged fifth maxillary tooth, prominent preorbital ridges, a large supraoccipital exposure on the skull table, a palatine-pterygoid suture anterior to the posterior end of the suborbital fenestra, and a pterygoid forming a neck surrounding the choana is unique to this species. Unlike previous phylogenies, our parsimony analysis recovers a monophyletic Late Cretaceous to Paleogene East to Southeastern Asian alligatoroid group, here named Orientalosuchina. The group includes Orientalosuchus naduongensis, Krabisuchus siamogallicus, Eoalligator chunyii, Jiangxisuchus nankangensis and Protoalligator huiningensis, all of them sharing a medial shifted quadrate foramen aerum. The recognition of this clade indicates at least two separate dispersal events from North America to Asia: one during the Late Cretaceous by Orientalosuchina and one by the ancestor of Alligator sinensis during the Paleogene or Neogene, the timing of which is poorly constrained.

A new Miocene ape and locomotion in the ancestor of great apes and humans.

Many ideas have been proposed to explain the origin of bipedalism in hominins and suspension in great apes (hominids); however, fossil evidence has been lacking. It has been suggested that bipedalism in hominins evolved from an ancestor that was a palmigrade quadruped (which would have moved similarly to living monkeys), or from a more suspensory quadruped (most similar to extant chimpanzees)1. Here we describe the fossil ape Danuvius guggenmosi (from the Allgäu region of Bavaria) for which complete limb bones are preserved, which provides evidence of a newly identified form of positional behaviour-extended limb clambering. The 11.62-million-year-old Danuvius is a great ape that is dentally most similar to Dryopithecus and other European late Miocene apes. With a broad thorax, long lumbar spine and extended hips and knees, as in bipeds, and elongated and fully extended forelimbs, as in all apes (hominoids), Danuvius combines the adaptations of bipeds and suspensory apes, and provides a model for the common ancestor of great apes and humans.

Possible species-flock scenario for the evolution of the cyprinid genus Capoeta (Cypriniformes: Cyprinidae) within late Neogene lake systems of the Armenian Highland.

We studied 4 Ma old isolated pharyngeal teeth from lake sediments of Çevirme (Tekman Palaeolake, Erzurum Province). Based on shape characters defined for 3D models of modern species, we found that the Pliocene lake constitutes sympatric occurrence of four Capoeta species (C. cf. umbla, C. cf. baliki, C. cf. sieboldi and C. sp. sevangi/capoeta), whose modern relatives belong to a monophyletic clade inhabiting today three different drainage systems of this region (Euphrates River, Kura River and Black Sea). We interpreted this high local diversity of closely related species in terms of the species-flock model. The Tekman palaeolake was a part of an unrecognized extended late Miocene to Pliocene palaeolake system in the present-day Armenian Highland, which has been disrupted by Pliocene tectonic activities. Surface uplift of the Armenian Highland contributed to the very characteristic biogeographic distribution and endemism of Capoeta in West Asian drainage systems. Thus, we proposed a species-flock scenario for the evolution and dispersal of the cyprinid genus Capoeta in a huge unrecognized palaeolake system in the present-day Armenian Highland.

A new testudinoid turtle from the middle to late Eocene of Vietnam.

BACKGROUND: Testudinoidea is a major clade of turtles that has colonized different ecological environments across the globe throughout the Tertiary. Aquatic testudinoids have a particularly rich fossil record in the Tertiary of the northern hemisphere, but little is known about the evolutionary history of the group, as the phylogenetic relationships of most fossils have not been established with confidence, in part due to high levels of homoplasy and polymorphism. METHODS: We here focus on describing a sample of 30 testudinoid shells, belonging to a single population that was collected from lake sediments from the middle to late Eocene (35-39 Ma) Na Duong Formation in Vietnam. The phylogenetic placement of this new material is investigated by integrating it and 11 other species of putative geoemydids from the Eocene and Oligocene to a recently published matrix of geoemydid turtles, that embraces the use of polymorphic characters, and then running a total-evidence analysis. RESULTS: The new material is highly polymorphic, but can be inferred with confidence to be a new taxon, Banhxeochelys trani gen. et sp. nov. It shares morphological similarities with other southeastern Asian testudinoids, Isometremys lacuna and Guangdongemys pingi, but is placed phylogenetically at the base of Pan-Testuguria when fossils are included in the analysis, or as a stem geoemydid when other fossils are deactivated from the matrix. The vast majority of other putative fossil geoemydids are placed at the base of Pan-Testuguria as well. DISCUSSION: The phylogenetic placement of fossil testudinoids used in the analysis is discussed individually and each species compared to Banhxeochelys trani gen. et sp. nov. The high levels of polymorphism observed in the new taxon is discussed in terms of ontogenetic and random variability. This is the first time that a large sample of fossil testudinoids has its morphological variation described in detail.

Earliest evidence of caries lesion in hominids reveal sugar-rich diet for a Middle Miocene dryopithecine from Europe.

The formation of dental caries is mainly caused by dietary habits and therefore, may contain information for dietary reconstructions of fossil hominids. This study investigates the caries lesion in the 12.5 Ma old type specimen of Dryopithecus carinthiacus Mottl 1957 (Primates, Hominidae) from St. Stefan (Austria). Potential food sources are identified on associated palynological data, which allow conclusions about food quality, sugar availability and the hominid metabolism during the Middle Miocene. Using micro computed tomography (µCT) and scanning electron microscopy (SEM) we provide a detailed analysis and characterization of the individuals' caries type. Its lesion is compared with a dataset of 311 wild chimpanzees, indicating morphological and etiological differences in caries formation between both species. The affected molar of D. carinthiacus reveals features known from severe dental caries in humans: (1) Cavitation with steep walls and smooth surface; (2) Reparative dentine at the roof of the pulp chamber; (3) Sclerotic dentine below the cavitation; (4) Association with dental calculus and (5) Unilateral usage of the healthy right tooth row. Its advanced primary caries, initiating on the intact enamel surface, indicates a frequent intake of highly cariogenic sugar-rich fruits, which likely exceeds the frugivory of extant chimpanzees. This finding corresponds with the associated palynological record, which infers a habitat with nearly year-round supply (9-10 months/year) of high quality foods (>carbohydrates; < fibers). Our conclusions challenge the model of a step-wise increase in dietary quality during hominid evolution and support the uricase hypothesis, which discusses the hominid autapomorphy of a fructose-based fat accumulation for periods of starvation. This model receives further validation by the identification of soft-tissue preservation, interpreted as fossilized white adipose cells, in the articulated hominid skeleton of Oreopithecus bamboli from Italy.

Potential hominin affinities of Graecopithecus from the Late Miocene of Europe.

The split of our own clade from the Panini is undocumented in the fossil record. To fill this gap we investigated the dentognathic morphology of Graecopithecus freybergi from Pyrgos Vassilissis (Greece) and cf. Graecopithecus sp. from Azmaka (Bulgaria), using new µCT and 3D reconstructions of the two known specimens. Pyrgos Vassilissis and Azmaka are currently dated to the early Messinian at 7.175 Ma and 7.24 Ma. Mainly based on its external preservation and the previously vague dating, Graecopithecus is often referred to as nomen dubium. The examination of its previously unknown dental root and pulp canal morphology confirms the taxonomic distinction from the significantly older northern Greek hominine Ouranopithecus. Furthermore, it shows features that point to a possible phylogenetic affinity with hominins. G. freybergi uniquely shares p4 partial root fusion and a possible canine root reduction with this tribe and therefore, provides intriguing evidence of what could be the oldest known hominin.

Messinian age and savannah environment of the possible hominin Graecopithecus from Europe.

Dating fossil hominids and reconstructing their environments is critically important for understanding human evolution. Here we date the potentially oldest hominin, Graecopithecus freybergi from Europe and constrain the environmental conditions under which it thrived. For the Graecopithecus-bearing Pikermi Formation of Attica/Greece, a saline aeolian dust deposit of North African (Sahara) provenance, we obtain an age of 7.37-7.11 Ma, which is coeval with a dramatic cooling in the Mediterranean region at the Tortonian-Messinian transition. Palaeobotanic proxies demonstrate C4-grass dominated wooded grassland-to-woodland habitats of a savannah biome for the Pikermi Formation. Faunal turnover at the Tortonian-Messinian transition led to the spread of new mammalian taxa along with Graecopithecus into Europe. The type mandible of G. freybergi from Pyrgos (7.175 Ma) and the single tooth (7.24 Ma) from Azmaka (Bulgaria) represent the first hominids of Messinian age from continental Europe. Our results suggest that major splits in the hominid family occurred outside Africa.

Neogene amphibians and reptiles (Caudata, Anura, Gekkota, Lacertilia, and Testudines) from the south of Western Siberia, Russia, and Northeastern Kazakhstan.

BACKGROUND: The present-day amphibian and reptile fauna of Western Siberia are the least diverse of the Palaearctic Realm, as a consequence of the unfavourable climatic conditions that predominate in this region. The origin and emergence of these herpetofaunal groups are poorly understood. Aside from the better-explored European Neogene localities yielding amphibian and reptile fossil remains, the Neogene herpetofauna of Western Asia is understudied. The few available data need critical reviews and new interpretations, taking into account the more recent records of the European herpetofauna. The comparison of this previous data with that of European fossil records would provide data on palaeobiogeographic affiliations of the region as well as on the origin and emergence of the present-day fauna of Western Siberia. An overview of the earliest occurrences of certain amphibian lineages is still needed. In addition, studies that address such knowledge gaps can be useful for molecular biologists in their calibration of molecular clocks. METHODS AND RESULTS: In this study, we considered critically reviewed available data from amphibian and reptile fauna from over 40 Western Siberian, Russian and Northeastern Kazakhstan localities, ranging from the Middle Miocene to Early Pleistocene. Herein, we provided new interpretations that arose from our assessment of the previously published and new data. More than 50 amphibians and reptile taxa were identified belonging to families Hynobiidae, Cryptobranchidae, Salamandridae, Palaeobatrachidae, Bombinatoridae, Pelobatidae, Hylidae, Bufonidae, Ranidae, Gekkonidae, Lacertidae, and Emydidae. Palaeobiogeographic analyses were performed for these groups and palaeoprecipitation values were estimated for 12 localities, using the bioclimatic analysis of herpetofaunal assemblages. CONCLUSION: The Neogene assemblage of Western Siberia was found to be dominated by groups of European affinities, such as Palaeobatrachidae, Bombina, Hyla, Bufo bufo, and a small part of this assemblage included Eastern Palaearctic taxa (e.g. Salamandrella, Tylototriton, Bufotes viridis). For several taxa (e.g. Mioproteus, Hyla, Bombina, Rana temporaria), the Western Siberian occurrences represented their most eastern Eurasian records. The most diverse collection of fossil remains was found in the Middle Miocene. Less diversity has been registered towards the Early Pleistocene, potentially due to the progressive cooling of the climate in the Northern Hemisphere. The results of our study showed higher-amplitude changes of precipitation development in Western Siberia from the Early Miocene to the Pliocene, than previously assumed.

Pronounced peramorphosis in lissamphibians--Aviturus exsecratus (Urodela, Cryptobranchidae) from the Paleocene-Eocene Thermal Maximum of Mongolia.

BACKGROUND: The oldest and largest member of giant salamanders (Cryptobranchidae) Aviturus exsecratus appears in the latest Paleocene (near the Paleocene-Eocene Thermal Maximum) of Mongolia. Based on femoral and vertebral morphology and metrics, a terrestrial adaptation has been supposed for this species. METHODOLOGY/PRINCIPAL FINDINGS: A detailed morphological reinvestigation of published as well as unpublished material reveals that this salamander shows a vomerine dentition that is posteriorly shifted and arranged in a zigzag pattern, a strongly developed olfactory region within the cranial cavity, and the highest bone ossification and relatively longest femur among all fossil and recent cryptobranchids. CONCLUSIONS/SIGNIFICANCE: The presence of these characteristics indicates a peramorphic developmental pattern for Aviturus exsecratus. Our results from Av. exsecratus indicate for the first time pronounced peramorphosis within a crown-group lissamphibian. Av. exsecratus represents a new developmental trajectory within both fossil and recent lissamphibian clades characterized by extended ontogeny and large body size, resembling the pattern known from late Paleozoic eryopines. Moreover, Av. exsecratus is not only a cryptobranchid with distinctive peramorphic characters, but also the first giant salamander with partially terrestrial (amphibious) lifestyle. The morphology of the vomers and dentaries suggests the ability of both underwater and terrestrial feeding.

The antiquity of the Rhine River: stratigraphic coverage of the dinotheriensande (eppelsheim formation) of the Mainz Basin (Germany).

BACKGROUND: Mammalian fossils from the Eppelsheim Formation (Dinotheriensande) have been a benchmark for Neogene vertebrate palaeontology since 200 years. Worldwide famous sites like Eppelsheim serve as key localities for biochronologic, palaeobiologic, environmental, and mammal community studies. So far the formation is considered to be of early Late Miocene age (~9.5 Ma, Vallesian), representing the oldest sediments of the Rhine River. The stratigraphic unity of the formation and of its fossil content was disputed at times, but persists unresolved. PRINCIPAL FINDINGS: Here we investigate a new fossil sample from Sprendlingen, composed by over 300 mammalian specimens and silicified wood. The mammals comprise entirely Middle Miocene species, like cervids Dicrocerus elegans, Paradicrocerus elegantulus, and deinotheres Deinotherium bavaricum and D. levius. A stratigraphic evaluation of Miocene Central European deer and deinothere species proof the stratigraphic inhomogenity of the sample, and suggest late Middle Miocene (~12.5 Ma) reworking of early Middle Miocene (~15 Ma) sediments. This results agree with taxonomic and palaeoclimatic analysis of plant fossils from above and within the mammalian assemblage. Based on the new fossil sample and published data three biochronologic levels within the Dinotheriensand fauna can be differentiated, corresponding to early Middle Miocene (late Orleanian to early Astaracian), late Middle Miocene (late Astaracian), and early Late Miocene (Vallesian) ages. CONCLUSIONS/SIGNIFICANCE: This study documents complex faunal mixing of classical Dinotheriensand fauna, covering at least six million years, during a time of low subsidence in the Mainz Basin and shifts back the origination of the Rhine River by some five million years. Our results have severe implications for biostratigraphy and palaeobiology of the Middle to Late Miocene. They suggest that turnover events may be obliterated and challenge the proposed 'supersaturated' biodiversity, caused by Middle Miocene superstites, of Vallesian ecosystems in Central Europe.

Bio-magnetostratigraphy and environment of the oldest Eurasian hominoid from the Early Miocene of Engelswies (Germany).

The paleobiogeography of hominoids exhibits a puzzling pattern of migrations between and within Africa and Eurasia. A precise dating of hominoid-bearing localities is therefore essential to reveal the timing, direction and possible causes of dispersals. Here, we present a bio-magnetostratigraphic analysis of the section of Engelswies (Southern Germany, Upper Freshwater Molasse, North Alpine Foreland Basin) where the oldest Eurasian hominoid was found. Our paleomagnetic results reveal a very short normal and a reverse magnetic polarity for the entire section. The polarity record is correlated to the Astronomical Tuned Neogene Time Scale using an integrated stratigraphic approach. This approach follows the chronostratigraphic framework for the Upper Freshwater Molasse, which combines magnetostratigraphy with biostratigraphic, lithostratigraphic and (40)Ar/(39)Ar dating results. According to this outcome, the reverse polarity of the Engelswies section most likely correlates to magnetochron C5Cr. The origin of the short normal polarity remains enigmatic. The magnetostratigraphic calibration and the evolutionary level of the Engelswies small mammal fauna suggest an age of 17.1-17.0Ma (Early Karpatian, Early Miocene) for the oldest Eurasian hominoid, and roughly confirm the estimates of Heizmann and Begun (2001). The estimated age suggests that the first hominoids in Eurasia are contemporaneous with Afro-Arabian afropithecins, and dispersal may have been facilitated by intra-Burdigalian (∼18-17Ma) sea-level low stands and the beginning of the Miocene Climate Optimum. The paleoclimatic and environmental reconstruction of the Engelswies locality indicates a lakeshore environment near dense subtropical rain forest vegetation, where paratropical temperatures (mean annual temperature around 20°C) and humid conditions (mean annual precipitation>1.100mm) prevailed.

Phylogenetic relationships and biogeographical patterns in Circum-Mediterranean subfamily Leuciscinae (Teleostei, Cyprinidae) inferred from both mitochondrial and nuclear data.

BACKGROUND: Leuciscinae is a subfamily belonging to the Cyprinidae fish family that is widely distributed in Circum-Mediterranean region. Many efforts have been carried out to deciphering the evolutionary history of this group. Thus, different biogeographical scenarios have tried to explain the colonization of Europe and Mediterranean area by cyprinids, such as the "north dispersal" or the "Lago Mare dispersal" models. Most recently, Pleistocene glaciations influenced the distribution of leuciscins, especially in North and Central Europe. Weighing up these biogeographical scenarios, this paper constitutes not only the first attempt at deciphering the mitochondrial and nuclear relationships of Mediterranean leuciscins but also a test of biogeographical hypotheses that could have determined the current distribution of Circum-Mediterranean leuciscins. RESULTS: A total of 4439 characters (mitochondrial + nuclear) from 321 individuals of 176 leuciscine species rendered a well-supported phylogeny, showing fourteen main lineages. Analyses of independent mitochondrial and nuclear markers supported the same main lineages, but basal relationships were not concordant. Moreover, some incongruence was found among independent mitochondrial and nuclear phylogenies. The monophyly of some poorly known genera such as Pseudophoxinus and Petroleuciscus was rejected. Representatives of both genera belong to different evolutionary lineages. Timing of cladogenetic events among the main leuciscine lineages was gained using mitochondrial and all genes data set. CONCLUSIONS: Adaptations to a predatory lifestyle or miniaturization have superimposed the morphology of some species. These species have been separated into different genera, which are not supported by a phylogenetic framework. Such is the case of the genera Pseudophoxinus and Petroleuciscus, which real taxonomy is not well known. The diversification of leuciscine lineages has been determined by intense vicariant events following the paleoclimatological and hydrogeological history of Mediterranean region. We propose different colonization models of Mediterranean region during the early Oligocene. Later vicariance events promoted Leuciscinae diversification during Oligocene and Miocene periods. Our data corroborate the presence of leuciscins in North Africa before the Messinian salinity crisis. Indeed, Messinian period appears as a stage of gradually Leuciscinae diversification. The rise of humidity at the beginning of the Pliocene promoted the colonization and posterior isolation of newly established freshwater populations. Finally, Pleistocene glaciations determined the current European distribution of some leuciscine species.