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Observed paleoenvironmental fluctuations in the North Alpine Foreland Basin, as one of the largest sedimentary archives of the Oligocene and Miocene are mainly controlled by regional factors. Global climate signals are usually less prominent than local tectonics and sedimentary input, caused by the enclosed paleogeographic setting of the Paratethys. Moreover are stratigraphic concepts still under debate, disabling a precise correlation of observed regional environmental changes to global climate patterns. In this study, a multi-proxy approach is used to achieve an accurate chronostratigraphy of regional formations and to verify whether global signals can be detected in the North Alpine Foreland Basin. Therefore, a detailed paleoenvironmental and biostratigraphic study of an 18 m-thick section of marine Miocene deposits (Neuhofen Formation) was carried out, using micropaleontology, sedimentology and geochemistry. In total 39 samples, yielding 68 foraminifera species and 47 ostracod species were processed together with 32 nannoplankton samples. Additionally, 34 ostracods and 49 benthic foraminifera were used for the analysis Oxygen and Carbon isotopes. Furthermore, 50 samples of six different sites in the Neuhofen Formation were used for statistical analyses of benthic foraminifera to assess supra-regional environmental correlations. Finally, the chronostratigraphic concept of the Neuhofen Formation was revised using magnetostratigraphic data from four sections, nannoplankton biostratigraphy and Sr-Isotope stratigraphy from previous studies as well as 3D-modelling using previous data additionally to 29 drillings. For the hypothesis that regional environmental patterns are correlating with global climate signals, environmental indices of the Neuhofen Formation (Isotopes, Diversity, Infaunalisation, Abundancy) were compared with global isotope values and Milankovic Cycles. The new stratigraphic concept of the Neuhofen Formation yielded an age of 18.1 – 17.6 Ma with a depositional time of 500,000 years. It was shown by a cluster analysis that strong faunal differences exist between the single localities, indicating separate paleoenvironments. These environmental differences are rather caused by regional factors. Occasionally, e.g. at 17.67 Ma, throughout the deposition of shallow marine sediments in the Neuhofen Formation the influence of global climate change can be inferred.

An integrated stratigraphic study has been performed of the Lower Miocene deposits of the Poljanska and Mala sections (North Croatian Basin), coupled with 40Ar/39Ar dating, geochemical and mineralogical study of tuffitic and silty sand beds. The sedimentary record evidences deposition influenced by volcaniclastic input. The presence of marine, brackish and non– marine microfossils indicates complex palaeoenvironment, palaeoecology, and palaeobiogeography. Four different units were recognized based on their fossil content and sedimentological data. At the base of the Poljanska section (Unit P1), tuffitic sediments bear common endemic brackish, and rare marine to freshwater and terrestrial microfossils. Radiometric 40Ar/39Ar dating obtained on volcanic glass shards from this layer provided an age of 22.2 ± 1.9 Ma correlating with the earliest Early Miocene and indicating the existence of older volcanic activity in the NCB pro tempore. The succession transitions into the brackish-water lacustrine environments (Unit P2). This unit has two presumed different climatic events: a maximum temperature event in the lower part of the interval and one interruption with lower temperatures recognized in the upper part of the interval. The co-occurrence of limnic and marine species in the uppermost part of the investigated sediments (Unit P3) indicates a marine inflow into the coastal lake. It marks the beginning of a salina-type lake formation in Poljanska. In the Mala section (Unit M1), biostratigraphically dated to the NN4 Zone, a transgressive sequence is documented through the presence of marine calcareous nannoplankton, planktonic and benthic foraminifera, and ostracods.

The middle Miocene (upper Serravallian, lower Volhynian) deposits at Karpov Yar near Naslavcea, northern Moldova, are among the few settings in which fossil fish are preserved with otoliths in situ. Here, we describe the new gobiid †Moldavigobius helenae gen. et sp. nov. from this locality. The taxon is characterized by small size (up to 34.2 mm SL), a compact body (body depth 17–21% SL), a fan-shaped caudal fin, large ctenoid scales (Moldavigobius gen. nov., together with its sagitta shape, suggest a relationship with Lesueurigobius Whitley, 1950, but its fan-shaped caudal fin and the unique sulcus contour of the otoliths preclude its attribution to that genus. In addition, we re-assign an otolith-based species previously described as Knipowitschia suavis Schwarzhans, 2014 as a second member of †Moldavigobius gen. nov. Accordingly, †Moldavigobius gen. nov. was represented by at least two species in the Serravallian of the Eastern Paratethys (†M. helenae gen. et sp. nov., †M. suavis nov. comb.). Moreover, †M. suavis is also known from the Serravallian ichthyofauna of the SE Mediterranean. †Moldavigobius gen. nov. thus demonstrates the key role of fossil skeletal material with otoliths in elucidating the ancient diversity of the Gobioidei.

Studies of otoliths suggest that Gobioidei, which are among the most species-rich groups of modern-day vertebrates, were prominent elements of late middle Miocene (early Sarmatian sensu lato) fish faunas in Europe and Western Asia. However, few complete skeletons have come to light. Here, we report an assemblage of six marine gobiid species, based on skeletons preserved with otoliths in situ, from the lower Volhynian (lower Sarmatian s.l.) of Karpov Yar, Naslavcea, northern Moldova (Eastern Paratethys). Previously only one of these species had been reported from the Central Paratethys, based on its otoliths alone. Five new species representing four new genera are described: †Katyagobius prikryli gen. et sp. nov., Pseudolesueurigobius manfredi gen. et sp. nov., †Sarmatigobius compactus gen. et sp. nov., †Yarigobius decoratus gen. et sp. nov., and †Y. naslavcensis gen. et sp. nov. All six species share the following set of characters, suggesting that they represent a monophyletic clade: 27–29 vertebrae (of which 10 are abdominal); spines of first dorsal fin distally filamentous; second dorsal fin with spine and 14–16 soft rays; anal fin with spine and 13–15 soft rays; caudal fin longish-to-lanceolate; otoliths (sagittae) with rounded, trapezoid-to-squarish shape. Their skeletal features suggest that they are closely related to Lesueurigobius Whitley, 1950, but the otoliths preserved in situ do not support such a classification. The new fossils most likely represent a stem lineage of the European Aphia lineage, and indicate that the diversity of gobiid lineages 12 million years ago differed clearly from that observed today.

Occurrence of the European extinct cordylid genus ‘Bavaricordylus’ from the early middle Miocene of Switzerland was recently questioned. Reanalysis of the Swiss material supports its assignment to the lacertid genus Janosikia, the presence of which is, therefore, indicated for the first time outside from southern Germany. Based on dentary morphology, it seems that two Janosikia morphotypes were present in the Miocene, one with a smooth and a second with a rough lateral surface of the dentary. The lacertid genus Janosikia seems to be confined to Central Europe, thriving there through at least the first half of the Miocene (from ca. 22 Ma to ca. 15.7 Ma).

Otolith morphology is a widely accepted tool for species identification in teleost fish, but whether this holds true for very small species remains to be explored. Here, the saccular otoliths of the cryptobenthic Mediterranean clingfish Gouania (Gobiesocidae) are described for the first time. The new data, although preliminary, indicate that otolith morphology and morphometry support the recognition of the recently differentiated five species of Gouania in the Mediterranean Sea. Furthermore, otoliths of phylogenetically closely related Gouania species resemble each other more than do those of the more distantly related species.

Fossil cichlids from East Africa offer unique insights into the evolutionary history and ancient diversity of the family on the African continent. Here we present three fossil species of the extinct haplotilapiine cichlid †Baringochromis gen. nov. from the upper Miocene of the palaeolake Waril in Central Kenya, based on the analysis of a total of 78 articulated skeletons. †Baringochromis senutae sp. nov., †B. sonyii sp. nov. and †B. tallamae sp. nov. are superficially similar, but differ from each other in oral-tooth dentition and morphometric characters related to the head, dorsal fin base and body depth. These findings indicate that they represent an ancient small species flock. Possible modern analogues of palaeolake Waril and its species flock are discussed. The three species of †Baringochromis may have begun to subdivide their initial habitat by trophic differentiation. Possible sources of food could have been plant remains and insects, as their fossilized remains are known from the same place where †Baringochromis was found.

The otoliths of three mudskipper species from the Persian Gulf that are characterised by different degrees of amphibious adaptation were examined using SEM imaging and otolith morphometry. Scartela...

Otolith shape variation in the Ornate goby, Istigobius ornatus (Teleostei: Gobiidae), collected along the intertidal coasts of the Persian Gulf and Oman Sea, was analysed using Wavelet transform technique in the ShapeR package to determine population differentiation and structure. There were significant differences (P 0.01). Mantel test showed a positive correlation between the Euclidean distance of otolith shape and geographical distances among populations (r = 0.93, P

Mudskippers are amphibious gobies (Teleostei: Gobioidei, Oxudercinae) that have served as models for the specialised physiology and behaviour of fishes out of water. In this study, a comparative analysis of the posterior vertebral column and the caudal skeleton of ten mudskipper species was conducted on the basis of X‐ray imaging. The species considered were Apocryptes bato, Apocryptodon madurensis, Boleophthalmus dussumieri, Oxuderces dentatus, Periophthalmodon freycineti, Pn. schlosseri, Periophthalmus novemradiatus, Ps. waltoni, Pseudapocryptes borneensis, and Scartelaos tenuis. For the osteological description the new term 'modified caudal vertebra' is used for all those vertebrae that display visibly modified neural and/or haemal spines compared to the spines of a 'usual' caudal vertebra, but are not involved in the support of caudal rays. The results reveal that the most terrestrial forms (Pn. freycineti, Pn. schlosseri, Ps. novemradiatus, Ps. waltoni) possess distinct traits that are seldom found in the other species. Among these features are (a) the existence of at least two modified caudal vertebrae (also present in S. tenuis), (b) a particularly close, dovetailing association between the neural spines of the preural vertebrae two and three (restricted to Ps. novemradiatus and Ps. waltoni), and (c) thickening and shortening of the ventralmost principal caudal rays (also present in B. dussumieri and S. tenuis). These findings support the idea that the posterior caudal vertebrae and caudal skeleton of the mentioned species are modified to enhance locomotion on land. Moreover, a relationship between character development and degree of terrestrial adaptation is probable, as all three traits are most pronounced in Ps. waltoni, which correlates with its strikingly high level of adaptation to amphibious life. A further aspect of this study is that the newly recognized skeletal structures have good fossilization potential and could therefore facilitate recognition of fossil species of mudskippers, which are currently unknown.

The Clupeiformes, including among others herrings, anchovies, shads and menhadens are ecologically and commercially important, yet their phylogenetic relationships are still controversial. Previous classification of Clupeiformes were based on morphological characters or lack of synapomorphic characters. More recent studies based on molecular data as well as new morphological evidence are keeping challenging their phylogenetic relations and there is still no consensus on many interrelationships within the Clupeiformes. In this study, we collected nuclear sequence data from 4,434 single-copy protein coding loci using a gene-capture method. We obtained a robust phylogeny based on 1,165 filtered loci with less than 30 % missing data. Our major findings include: 1) reconfirmation of monophyly of the Clupeiformes, that is, Denticipitidae is sister to all other clupeiforms; 2) the polyphyletic nature of dussumieriids and early branching of Spratelloididae from all other clupeoids were confirmed using datasets curated for less missing data and more balanced base composition in the respective taxa. The next branching clade is the monophyletic Engraulidae. Pristigasteridae also is monophyletic, but it was nested in the previously defined "Clupeidae". Within Pristigasteridae there is no support for monophyletic Pelloninae. Chirocentrus is close to Dussumieria and not to engraulids. The miniaturized Sundasalanx is placed close to the ehiravine Clupeonella, however, with a relatively deep split. The genus Clupea, is not part of the diverse "Clupeidae", but part of a clade containing additionally Sprattus and Etrumeus. Within the crown group clades, Alosidae and Dorosomatidae are retrieved as sister clades. Based on new fossil calibration points, we found that major lineages of the clupeiforms diverged in the late Cretaceous and early Paleogene. The extinction event at the end of the Cretaceous may have created ecological niches, which could have fueled the diversification of clupeiform fishes. Based on the strong evidence of the present study, we propose an updated classification of Clupeiformes consisting of ten families: Denticipitidae; Spratelloididae; Engraulidae (Engraulinae + Coiliinae); Clupeidae; Chirocentridae; Dussumieriidae; Pristigasteridae; Ehiravidae; Alosidae, Dorosomatidae.

The suborder Gobioidei is among the most diverse groups of vertebrates, comprising about 2310 species. In the fossil record gobioids date back to the early Eocene (c. 50 m.y. ago), and a considerable increase in numbers of described species is evident since the middle Miocene (c. 16 m.y. ago). About 40 skeleton-based gobioid species and > 100 otolith-based species have been described until to date. However, assignment of a fossil gobioid species to specific families has often remained tentative, even if well preserved complete specimens are available. The reasons are that synapomorphies that can be recognized in a fossil skeleton are rare (or absent) and that no phylogenetic framework applicable to gobioid fossils exists. Here we aim to overcome this problem by developing a phylogenetic total evidence framework that is suitable to place a fossil skeleton-based gobioid at family level. Using both literature and newly collected data we assembled a morphological character matrix (48 characters) for 29 extant species, representing all extant gobioid families, and ten fossil gobioid species, and we compiled a multi-gene concatenated alignment (supermatrix; 6271 bp) of published molecular sequence data for the extant species. Bayesian and Maximum Parsimony analyses revealed that our selection of extant species was sufficient to achieve a molecular ‘backbone’ that fully conforms to previous molecular work. Our data revealed that inclusion of all fossil species simultaneously produced very poorly resolved trees, even for some extant taxa. In contrast, addition of a single fossil species to the total evidence data set of the extant species provided new insight in its possible placement at family level, especially in a Bayesian framework. Five out of the ten fossil species were recovered in the same family as had been suggested in previous works based on comparative morphology. The remaining five fossil species had hitherto been left as family incertae sedis. Now, based on our phylogenetic framework, new and mostly well supported hypotheses to which clades they could belong can be presented. We conclude that the total evidence framework presented here will be beneficial for all future work dealing with the phylogenetic placement of a fossil skeleton-based gobioid and thus will help to improve our understanding of the evolutionary history of these fascinating fishes. Moreover, our data highlight that increased sampling of fossil taxa in a total-evidence context is not universally beneficial, as might be expected, but strongly depends on the study group and peculiarities of the morphological data.

The marine middle Burdigalian sediments of the North Alpine Foreland Basin (NAFB) or Molasse Basin are well known for abundant occurrences of benthic foraminifers and ostracods. However, taxonomic studies of ostracod assemblages are comparatively rare and, given its abundance, the group remains heavily understudied. Here we report a new, rich and well-preserved ostracod fauna from the middle Burdigalian Neuhofen Formation at Mitterdorf, in the northeastern part of the NAFB. The material comprises a total of 3029 ostracod specimens. We identified 44 species; all species are described and, with one exception, illustrated. Our material reveals the presence of 17 ostracod species that were previously unknown from the middle Burdigalian in the NAFB, i.e., Cytherella aff. vulgatella, Cytherella aff. mehesi, Bairdoppilata aff. subdeltoidea, Paranesidea? sp., Paracypris aff. aerodynamica, Parakrithe dactylomorpha, Pseudopsammocythere sp., Callistocythere daedalea, Costa aff. reticulata, Ruggieria aff. longecarenata, Heliocythere aff. vejhonensis, Heliocythere sp., Loxocorniculum hastatum, Neocytherideis cypria, Cytheretta cf. tenuipunctata dentata, Cytheropteron cf. ruggierii, and Semicytherura resecta. The palaeobiogeography of the ostracods indicates a relatively open marine Burdigalian connection between the NAFB and the Mediterranean Sea, in agreement with previous work. Our data also suggest that a significant degree of endemism existed in the middle Burdigalian sea of the NAFB, of which previous studies yielded only sparse signs. Moreover, our results indicate that some ostracod species survived the late Burdigalian sea retreat from the NAFB by dispersing eastward into the adjacent regions (Vienna Basin, Carpathian Foredeep), where marine conditions still prevailed. A compilation of all known middle Burdigalian ostracod faunas from the NAFB documents a high species diversity, with over 120 species. Their spatial distribution, with many species being restricted to a single site, suggests heterogenous microhabitats, possibly influenced by small-scale differences in water depth, nutrient availability and oxygenation.

Otolith morphology in the tooth-carp/killifish genus Aphanius is an important source of taxonomic information. However, little is known about the range of ontogenetic variation in otolith morphology. In this study, the development of otolith morphology during the early life stages of the Zagros tooth-carp, Aphanius vladykovi is described and discussed. The study material comprises 15 larvae and early juveniles representing nine different developmental stages (0–120 days post hatching), all reared under the same laboratory conditions. The results reveal (i) clear ontogenetic differences in otolith morphology during larval and early juvenile growth phases, (ii) a developmental progression in otolith morphology, which is similar to that described for A. farsicus in a previous study, and (iii) a more rapid rate of otolith development during the larval phase in A. vladykovi than in A. farsicus. It can be hypothesized that the shared pattern of overall development seen in A. vladykovi and A. farsicus reflects their close phylogenetic relationship.

The taxonomic validity of the recently described endemic species Aphaniops hormuzensis (Cyprinodontiformes: Aphaniidae) from southern Iran has been questioned as its original description did not include data from another recently described endemic species, i.e. A. kruppi from eastern Oman. Because of the absence of mitochondrial COI gene sequences, Aphaniops hormuzensis was also not included in recent phylogenetic studies. We therefore generated COI gene data for A. hormuzensis and used the new data to construct a phylogenetic tree that also included A. kruppi (amongst other species). Bayesian and Maximum likelihood phylogenetic analyses reveal that both A. hormuzensis and A. kruppi are monophyletic, with high support values. We conclude that A. hormuzensis is a valid species, which is distinct from A. kruppi and also from A. ginaonis, another tooth-carp endemic to southern Iran. Our results add further support to the taxonomic distinctness of A. hormuzensis from southern Iran.

The Pannonian Basin of Croatia is the largest back-arc extensional basin on the European continent, located between the Alps, Carpathians and Dinarides. Syn-rift subsidence started at ~18 Ma and predated the onset of the Miocene Climate Optimum (MCO; 17–14.7 Ma). In this paper, we investigate the evolution of the fluvial-lacustrine palaeoenvironment just prior to the transgression of the Central Paratethys Sea. Studies are based on a 17-m-thick section of the Požega palaeolake in the Southern Pannonian Basin using palaeobiological (mollusks, ostracods, palynomorphs, fish otoliths), mineralogical and sedimentological data. Our results reveal fluctuating freshwater and brackish conditions and a pulsating water budget, which we relate to phases in which the basin was, alternatively, hydrologically open and closed. We infer that the alternating environmental conditions can be best-explained by dry and wet periods, respectively. Droughts produced stagnant conditions, algae overproduction and increased salinity levels, supporting the survival of sporadically invading brackish water species. The wet periods, on the other hand, led to lake highstand episodes, expansion of lake area, and to the widespread development of profundal facies. The lake basin eventually became overfilled and connected through outflowing rivers with the Central Paratethys sea. However, the absence of brackish-marine assemblages in the Požega palaeolake nevertheless indicates that there was no marine water inflow from the Central Paratethys at that time.

The regional and stratigraphic subdivision of the Tertiary as part of the Stratigraphic Table of Germany 2016 (STD 2016) is summarised. Primarily, changes compared to STD 2002 are commented. Concerning more detailed explanations, the reader is referred to the respective literature and corresponding definitions in LithoLex (https://litholex.bgr.de). The STD 2016 has not only practical significance, but is also a working basis for (1) the further development of the stratigraphic terminology of the German Tertiary, (2) the correlations of regional units and (3) an improved correlation of the regional stages of the North Sea Basin and the Central Paratethys with the Global Stratigraphic Scale.

The fossil record of cichlids is sparse, and every new discovery can provide new insights into the evolutionary history of this speciose freshwater fish family. In this article, we describe †Rebekkachromis gen. nov. from the middle-to-late Miocene (c. 11 MYA) of the Central Kenya Rift within the East African Rift system. †Rebekkachromis is represented by two species that differ from all other fossil and extant African cichlids, except Etia, in possessing the unique character combination of two supraneural bones and a set of robust tricuspid oral teeth in the outer row of the dentition. Furthermore, †Rebekkachromis exhibits the only proposed morphological synapomorphy of the Haplotilapiini, namely the presence of tricuspid teeth in the inner row of the oral dentition. We show that †Rebekkachromis constitutes the oldest reliably identified fossil record of a haplotilapiine. The evolution of cichlid fishes possessing tricuspid teeth in the rivers and lakes of the Central Kenya Rift during the middle-to-late Miocene could have been facilitated by volcanic activity, as repeated ash falls may well have fostered the growth of algae and in particular diatoms. These fishes could thus have had a major advantage, because they could exploit the newly available, rich food resources.

Otolith morphology in the tooth-carp/killifish genus Aphanius is a source of informative taxonomic characters at both the species and population level. Most work on otoliths has focused on adult specimens, while evidence of ontogenetic variation is rarely provided. In this study we describe the development of otolith morphology during the early life stages of an endangered and endemic species, the Fars tooth-carp Aphanius farsicus from southern Iran. The study material comprises 34 larvae and early juveniles representing nine different developmental stages (0–120 days post hatching), all reared under the same laboratory conditions. The results reveal (i) a significant correlation between standard length and otolith size (length) in larval and early juvenile stages, (ii) clear differences in otolith morphology between larvae/early juveniles and adults, and (iii) a temporal link between the appearance of the sulcus on the otolith’s inner face and the emergence of the dorsal and anal fins. Our results indicate that otoliths of Aphanius can be recognized as originating from larval or early juvenile fish based on their short rostrum and antirostrum lengths and wide excisura, in addition to their small size. These immature otoliths are, however, not diagnostic at the species level in A. farsicus, nor most probably in other species of tooth-carp. The outcome of our study is also of interest to palaeontologists working with fossil killifish otoliths, as it can help avoid misinterpretation of ancient species diversity.

For Oligocene and Miocene sediments of the Central Paratethys realm, regional chronostratigraphic stages have been defined on the basis of characteristic faunal assemblages, often containing abundant endemic elements. Although considerable progress has been made in correlating Paratethyan stages with the Global Time Scale (GTS), the task remains incomplete, especially for the Early Miocene period. The present study focuses on Lower Miocene deposits of the Central Paratethys in southeastern Germany, i.e. the Upper Marine Molasse (OMM) and Upper Brackish Molasse (OBM) of the North Alpine Foreland Basin (NAFB, Molasse Basin). The sediments concerned are assigned to the regional Ottnangian stage of the Central Paratethys, which corresponds to the middle Burdigalian in the GTS. We present a formal lithostratigraphic definition of the Untersimbach and Neuhofen Formations (both OMM), combined with litho-, bio- and magnetostratigraphic investigations, which together provide an improved chronology for the middle Burdigalian interval in the NAFB. The base of the Ottnangian stage is shown to correlate with polarity chron C5En, and we consider an absolute age of around 18.2 Ma to be most plausible. As the OMM in the southeast German part of the NAFB represents a single transgressive-regressive sequence, our new data suggest that the Ottnangian transgression began during chron C5En (similar to 18.2 Ma), whereas the onset of the regressive phase lies within chron C5Dr.2r (similar to 18 Ma). The uppermost marine deposits (top OMM/"Glaukonitsande & Blattermergel") and the lowermost brackish sediments (OBM/Oncophora Formation) are correlated with polarity chron C5Dn (similar to 17.4 Ma). Moreover, our results clearly demonstrate that the lower Oncophora Formation (normal polarity) is not time-equivalent to the OBM/Kirchberg Formation in the southwest German sector of the NAFB (largely reverse polarity) as has hitherto been assumed. The lower Oncophora Formation is most probably contemporaneous with the lowermost OBM sediments (Grimmelfingen Formation) in the southwest German Molasse Basin.

Among the species of Aphanius Nardo, 1827, Aphanius dispar (Ruppell, 1828) is the most common taxon and has long been viewed as representing a species group rather than a single species. This study provides comprehensive data on the phylogenetic relationships, morphology, and otoliths within the A. dispar species group, including the description of a new species. Our data demonstrate that the "true" A. dispar is restricted to the Red Sea drainages and that all other populations hitherto identified as A. dispar actually represent separate species. Four main clades are defined and named for the geographic areas in which the respective species of Aphanius occur. The oldest one is the "Red Sea clade," it comprises A. dispar. The "Dead Sea clade" is represented by A. richardsoni (Boulenger, 1907). It is sister to both the "Hormuzgan clade" in S Iran (containing A. hormuzensis sp. nov. and A. ginaonis (Holly, 1929)) and the "Persian Gulf & Gulf of Oman clade" (comprising A. stoliczkanus (Day, 1872)). The species separation within the A. dispar group is confirmed by the distinctive otolith morphology of each species. Moreover, we present a time-calibrated phylogeny (chronogram) for the A. dispar species group using dagger A. princeps (16-17 Mya) as a minimum age and the first appearance of dagger Prolebias (33-34 Mya) as a maximum age for the genus Aphanius. The evolution and historical biogeography routes are discussed based on the outcome of the chronogram and in the context of the geological and climatic history of the Near East in Pliocene-Pleistocene times.

The present study was conducted to provide a comparative description of the caudal skeleton of the mullid fishes (family Mullidae) from the western Indo-Pacific region. Nine species belonging to three genera were studied, i.e., Mulloidichthys vanicolensis, Parupeneus heptacanthus, P. macronemus, P. margaritatus, P. rubescens, Upeneus doriae, U. sundaicus, U. tragula, and U. vittatus. Based on X-ray imaging, the configuration of the caudal skeleton and the caudal-fin formula are reported for all species. The new data reveal that the shape, size and degree of fusion of individual bones of the caudal skeleton and the caudal fin-ray formula can be different among genera, and also between congeneric species. The neural spine of PU2 is short in all species, except in P. macronemus. The hypural plates 1 and 2 are usually separated, but fused in M. vanicolensis and U. doriae. The hypural plates 3 and 4 are fused in all species except in P. rubescens. The parhypural is long and broad in all examined species, the highest relative length of the parhypural is present in P. macronemus. The uroneural 2 is absent in M. vanicolensis, U. doriae, U. tragula, and U. vittatus, but present in the remaining species. These differences indicate that the structures of the caudal skeleton are additional useful tools in the taxonomic study of mullid species.

Precise age constraints are rare for the Oligocene to Miocene sediments in the North Alpine Foreland Basin, also referred to as Molasse Basin. This is mainly due to the lack of index fossils, absence of continuous outcrops and scarcity of radiometrically datable rocks. In this study, we applied the Sr isotope dating technique to fossil teleost fish otoliths to determine absolute ages for Lower Miocene (mid-Burdigalian, lower Ottnangian) marine sediments. The fossil otoliths from the marine genera Diaphus and Coelorinchus used in our analysis were recovered from the Neuhofen Beds in the SE German Molasse Basin. For the first time, leaching experiments were conducted on different otolith fractions to extract the ambient Sr-87/(86) Sr seawater signal. Based on the latest version of the evolution curve for the Sr-87/(86) Sr ratio in the global ocean, our new data suggest an age of 17.1 +/- 0.3 Ma, which is at least 0.6 m. y. younger than hitherto assumed for the Neuhofen Beds based on biostratigraphy and lithostratigraphic correlations. We suggest that strong riverine inputs could account for this shift in the Sr isotope signal to younger ages.

Gobiidae (Gobiiformes, Teleostei) is among the largest families of vertebrates. These fishes are distributed worldwide and contribute significantly to species diversity in marine habitats and reefs. However, their fossil record is sparse prior to the Miocene and little is known about the course of diversification of the clade. Here we report exceptionally well-preserved skeletal remains of the oldest known Gobius from an Early Miocene (Burdigalian) marine ecosystem of Central Europe (Czech Republic). Gobius jarosi Přikryl & Reichenbacher sp. nov. is dated to 19.1–20.4 Ma by biostratigraphical analysis of calcareous nannoplankton from small fragments of the holotype matrix. Gobius jarosi sp. nov. is characterized by a pterygiophore formula of 3-22110 and a premaxilla with a distinctive postmaxillary process, has 11 abdominal and 16–17 caudal vertebrae, six thin spines in the first dorsal fin and one spine and 12 soft rays in the second dorsal fin, one spine and 11 rays in the anal fin, and two anal fin pte...

The modern Gobioidei (Teleostei) comprise eight families, but the extinct †Pirskeniidae from the lower Oligocene of the Czech Republic indicate that further families may have existed in the past. However, the validity of the †Pirskeniidae has been questioned and its single genus †Pirskenius has been assigned to the extant family Eleotridae in previous works. The objective of this study is to clarify the status of the †Pirskeniidae. Whether or not the †Pirskeniidae should be synonymised with the Eleotridae is also interesting from a biogeographical point of view as Eleotridae is not present in Europe or the Mediterranean Sea today. We present new specimens and re-examine the material on which the two known species of †Pirskenius are based (†P. diatomaceus Obrhelova, 1961; †P. radoni Přikryl, 2014). To provide a context for phylogenetically informative characters related to the palatine and the branchiostegal rays, three early-branching gobioids (Rhyacichthys, Protogobius, Perccottus), an eleotrid (Eleotris) and a gobiid (Gobius) were subjected to micro-CT analysis. The new data justify revalidation of the family †Pirskeniidae, and a revised diagnosis is presented for both †Pirskenius and †Pirskeniidae. Moreover, we provide for the first time an attempt to relate a fossil gobioid to extant taxa based on phylogenetic analysis. The results indicate a sister-group relationship of †Pirskeniidae to the Thalasseleotrididae + Gobiidae + Oxudercidae clade. Considering the fossil record, the arrival of gobioids in freshwater habitats in the early Oligocene apparently had generated new lineages that finally were not successful and became extinct shortly after they had diverged. There is currently no evidence that the Eleotridae was present in the European ichthyofauna in the past.

The African Cichlidae Oreochromis (Alcolapia) and Oreochromis amphimelas can survive in extremely alkaline environments and represent the only known modern alkaliphilic cichlid fish found in Africa. The presence of fossil cichlids from the Miocene of central Kenya (Tugen Hills) that are morphologically similar to Oreochromis (Alcolapia) has been noted in previous works, but the conclusions remained tentative. The purpose of this study is to examine newly discovered fossil cichlids from the Tugen Hills and to compare their osteology with that of extant Oreochromis (Alcolapia). This is performed based on a comprehensive collection of comparative material, using microscopy and computed microtomography (μCT). We provide a revised diagnosis for the genus †Rebekkachromis, and revise its systematic relationships by assigning it to the Oreochromini (rather than to the Etiini). Two new species of †Rebekkachromis are described, i.e., †R. valyricus, sp. nov., and †R. vancouveringae, sp. nov., and a morphologically diverse assemblage of co-occurring †Rebekkachromis specimens is documented. Moreover, we found that †Rebekkachromis had three sensory canal pores (instead of four) on the lower arm of the preopercle, a feature that distinguishes both the modern Oreochromis (Alcolapia) and our fossil specimens from almost all other modern African cichlid fish. Our new data indicate that alkaliphile cichlids similar to Oreochromis (Alcolapia) were present in Central Kenya about 10–13 Ma ago and that the ability of African cichlid fishes to thrive in highly alkaline waters had already developed by that time.

Gobies (Gobiidae + Oxudercidae) are among the largest groups of extant marine fishes. Fossils of gobies are abundant since the Miocene, and many species have been reported so far. However, delimitation of fossil goby species is challenging because molecular markers and diagnostic traits such as the disposition of sensory head papillae are lost. This study provides, for the first time, an actualistic framework for the identification of fossil goby species. We focus on characters that can in principle be recognized in fossils, and evaluate their ability to discriminate between extant goby species based on statistical analyses. Using 14 extant species of Gobius and seven species of Pomatoschistus, we conducted otolith morphometry, elliptic Fourier shape analysis of otoliths using the package ‘Momocs’, conventional fish morphometry, and meristic counts. In addition, the otoliths of all species are depicted based on SEM images and briefly described. Otolith Fourier shape analysis proved to be most efficient in discrimination of species within both genera, Gobius and Pomatoschistus. Several characters used in the other approaches also worked well, but the results were variable, and the relative taxonomic significance of particular variables tended to change depending on the species under consideration. We propose otolith shape analysis as a powerful tool to explore ancient goby species diversity when samples with abundant fossil otoliths are present. Overall, the herein presented data will greatly facilitate delimitation of fossil goby species in future studies, and will consequently shed new light on the evolution of goby diversity and biogeography through time.

A new genus and species of fossil cichlid fishes of middle Miocene age (12.5 Ma) is described from the Ngorora fish Lagerstätte (Tugen Hills, Kenya) in the East African Rift Valley. Parsimony analysis of morphological characters using published phylogenetic frameworks for extant cichlids combined with the application of a comprehensive best-fit approach based on morphology was employed to place the new fossil taxon in the phylogenetic context of the African cichlids. The data reveal that the fossil specimens can be assigned to the tribe Oreochromini within the haplotilapiines. †Oreochromimos kabchorensis gen. et sp. nov. shows a mosaic set of characters bearing many similarities to the almost pan-African Oreochromis and the East African lake-endemic Alcolapia. As the striking diversity of present-day African cichlids, with 1100 recognised species, has remained largely invisible in the fossil record, the material described here adds significantly to our knowledge of the Miocene diversity of the group. It effectively doubles the age of a fossil calibration point, which has hitherto been used to calibrate divergence times of the East African cichlids in molecular phylogenetic investigations. Furthermore, the comparative dataset derived from extant cichlids presented here will greatly facilitate the classification of fossil cichlids in future studies.

The North Alpine Foreland Basin (NAFB) experienced rapidly changing palaeoenvironmental conditions at the end of the Early Miocene (middle-to-late Burdigalian, Paratethys stages Ottnangian and Karpatian), with transformation of fully marine settings into brackish and eventually freshwater environments. These changes were related to global sea-level fluctuations, climate oscillations, and tectonic processes associated with the uplift of the Alps. This study presents a new and comprehensive data set, derived from 13 boreholes and 491 core samples from the Molasse Basin of southwest Germany, that provides a significantly better understanding of late Early Miocene palaeoenvironments in the NAFB. Based on lithostratigraphy, biostratigraphy, and palaeoecological interpretation of rich fauna and flora assemblages, we show that the upper Burdigalian sediments comprise a regressive lower segment (Grimmelfingen Formation and equivalents), as indicated by the occurrence of the bivalve Rzehakia, and a transgressive upper component (Kirchberg Formation and equivalents) characterized by the advent of new fish and mollusc assemblages. Furthermore, we present a new lithostratigraphic concept for the Kirchberg Formation, which forms an important part of the Upper Brackish Molasse (OBM). Based on biostratigraphic constraints and recently published magnetostratigraphic data, it appears that the regressive segment of the upper Burdigalian OBM sediments correlates with the end of the global third-order sea-level cycle Bur 3, while the transgressive component reflects the Karpatian transgression at the beginning of sea-level cycle Bur 4. This implies that the Karpatian transgression actually affected the Molasse Basin of southern Germany and Upper Austria, whereas previous studies had considered this part of the NAFB as terrestrial during that time span. The new results are depicted in three palaeogeographic maps for the NAFB at 18 Ma, 17.5–17.1 Ma and 17 Ma, respectively.

Studies of the taxonomy and present-day distribution of landlocked populations of the killifish Aphanius Nardo, 1827 (Cyprinodontidae) provide a key to understanding their zoogeographic history, and shape conservation strategies and habitat management. Here we report for the first time on the sympatric occurrence of the rare Mesopotamian tooth-carp A. mesopotamicus with a member of the widely distributed common tooth-carp (A. dispar group). Both were found in the Shadegan Wetland of the Mesopotamian drainage system (Southwestern Iran). External characters of individuals, otolith morphology and molecular data based on the cytochrome b gene confirm species identification. The otolith morphology of A. mesopotamicus, hitherto unknown, is strikingly similar to previously described otoliths of A. cf. pluristriatus from the Khonj stream (Southern Iran), which apparently reflects a close relationship between these recently diverged species. Several of the specimens recovered from the Shadegan Wetland showed intermediate characters in pigmentation, otolith morphology and some morphometric values. These specimens are interpreted as putative hybrids, which is additionally supported by a multivariate analysis of the morphometric data. Previous reports suggest that natural hybridization between species of Aphanius results largely from range overlap and range extension. A virtual distribution map derived from climatic modelling studies based on DIVA-GIS (7.5.0) indicates that sympatry of the A. dispar group and A. mesopotamicus is primarily determined by levels of precipitation during the warmest quarter, and is likely to occur only in Southwestern Iran.