Bettina Reichenbacher

Ludwig-Maximilians-Universität in Munich | LMU

Research interest in the diversity and evolutionary history of herring-like fossils (subcohort Clupei) has increased in recent decades. However, little is known about the relationships between fossils assigned to Clupei, particularly those that are demonstrably related to extant herring-like members of the order Clupeiformes. To help bridge this ga...

The killifish genus Aphaniops consists of nine species distributed in Eastern Africa and the Middle East. However, distinguishing these species from each other based on morphological traits is challenging. Here we investigate the utility of otoliths (sagittae) in distinguishing between A. dispar, A. ginaonis, A. hormuzensis, A. kruppi and A. stolic...

The extant gobioid fishes form a highly diverse group comprised of eight families and over 2000 species. They pose many taxonomic and phylogenetic challenges, particularly when working with fossils. Here we introduce †Simpsonigobius gen. nov., a new freshwater gobioid of small size (34 mm) from the Lower Miocene of Turkey, and analyse its relations...

To evaluate scale morphological variability (shape, size, topological macro‐ and microstructures, ornamentation patterns) of cyprinid fishes, nine species of the genus Garra were selected and their scales from five different body regions were studied by using light and scanning electron microscopy. The scales of the examined species were thin with...

The Lower Miocene (Aquitanian to ?lower Burdigalian) type locality of the Wiesbaden Formation in the northern region of the Mainz Basin (abandoned ‘Kalkofen Quarry’, Wiesbaden, State of Hesse, Germany) is known to reveal a regressive-dominated sequence encompassing marine, brackish to terrestrial sediments. Our documentation of this depositional se...

The high diversity of extant gobiids (Gobiidae: Teleostei) makes taxonomic and phylogenetic interpretation of fossil members of the clade a difficult task. To facilitate future taxonomic and systematic work on the group, we have assembled a morphological reference database encompassing skeletal characters, an otolith atlas and otolith morphometric...

The Miocene Climate Optimum (MCO) and the subsequent Miocene Climate Transition (MCT) are important biotic, environmental and geologic events. Here we address whether this holds true for the epicontinental Eastern Paratethys Sea (today's Black – Caspian Sea region). Two Tarkhanian – lower Chokrakian sequences of Middle Miocene age in the Kerch Peni...

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The South Caspian Sea sub-basin, owing to its complex paleogeographic history and habitat diversity, represents a center of endemism and a high-priority conservation area for the gobiid genus Ponticola. However, very little is currently known about most biological aspects of these species in general, and this sub-basin is highly threatened by anthr...

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 S...

Sex dimorphism is found in many organisms and is an important source of intraspecific variation. Among freshwater fishes, all members of the Aphaniidae (killifishes, Cyprinodontiformes) are known for their pronounced sex dimorphism, and it has been proposed that sexual selection has played a role in their diversification. However, few studies have...

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 otoli...

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 a...

There is a significant gap in our knowledge of the intraspecific morphological variability in freshwater fish, although such data are crucial for understanding species diversity. Here we use the killifish Aphaniops stoliczkanus (Day, 1872; Aphaniidae: Cyprinodontiformes), which is a widespread but poorly known freshwater species in the Middle East,...

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. sun...

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,...

Auch nach mehreren Jahrzehnten der Erforschung wartet die egerische Transgressionsabfolge von Unterrudling bei Eferding (Oberösterreich), welche den Übergang der Flachwasserablagerungen der Linz-Melk Formation zu den Tiefwassersedimenten der Eferding Formation (EF) umfasst, noch mit Überraschungen auf. So war kurzzeitig eine Schicht in der siltig-t...

Much remains to be learned about the past diversity and evolutionary history of the Clupeidae (herrings, shads and allies), owing to the frequently subtle differences between modern taxa and the moderate preservational quality of some fossils. In this study, new clupeid fossils are described from a new locality from the upper Miocene of the Serres...

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 7...

This article is part I of our study on a highly diverse assemblage of goby species from the lower Volhynian (lower Sarmatian sensu lato) deposits of northern Moldova (Karpov Yar, Naslavcea, western sector of the Eastern Paratethys). Six species, including five new ones, representing four new genera are described. All share the following unique feat...

The North Alpine Foreland Basin (NAFB) comprises one of the most complete sedimentary records of the Oligocene and Miocene. Driven by global sea-level fluctuations, vast sedimentary influx and tectonic movement. The locality of Unterrudling near Eferding (Upper Austria) exposes the largest succession of sedimentary deposits from the late Oligocene...

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 morpholog...

The Persian Gulf and Oman Sea are characterized by an interesting paleoclimatic history and different ecological settings, and offer a unique study area to investigate the genetic structure of marine organisms including fishes. The Ornate goby Istigobius ornatus is widely distributed throughout the tropical Indo-West Pacific including the Persian G...

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 otolit...

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 < 0.001, ANOVA) among geograph...

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 no...

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 previ...

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. Scartelaos tenuis, which is the most aquatic of the three species, has rhomboid to rectangular otoliths that conform to the general otolith type of many marine g...

Aphaniops dispar, widespread around the Arabian Peninsula, was recently separated in four species (A. dispar, A. hormuzensis, A. kruppi, A. stoliczkanus) by molecular results and colour patterns, but the morphological differences are small and call for more studies. Here we report differences in skeleton and median fin osteology of these species. I...

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 consi...

Background: The diversification process known as the Lake Tanganyika Radiation has given rise to the most speciose clade of African cichlids. Almost all cichlid species found in the lakes Tanganyika, Malawi and Victoria, comprising a total of 12-16 tribes, belong to this clade. Strikingly, all the species in the latter two lakes are members of the...

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. Thi...

Precise age data are a basic prerequisite for the correlation of a sedimentary succession with the Global Time Scale, which in turn allows one to place its biotic and other data in a global context. In the North Alpine Foreland Basin (NAFB), a patchy distribution of outcrops and uncertainties in the correlation of strata have led to conflicting age...

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 app...

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 w...

Otolith morphology in the tooth-carp/killifish genus Aphanius is an important source of taxo-nomic 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 discu...

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 represen...

The taxonomic information inscribed in otoliths has been widely ignored in ichthyological research, especially in descriptions of new fish species. One reason for this is that otolith descriptions are per se qualitative, and only a few studies have presented quantitative data that can support assignments of otoliths to individual species or permit...

Mudskippers are amphibious gobies (Teleostei: Gobioidei) that have served as models for the specialised physiology and behaviour of fishes out of water. Here we present new data on the skeletons and otoliths of three endemic mudskipper species from the Persian Gulf, i.e. Boleophthalmus dussumieri, Scartelaos tenuis and Periophthalmus waltoni, suppl...

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 palaeoenvironmen...

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 lif...

Based on a total of 161 otoliths, a unique teleost fauna is described from the middle Miocene in the eastern Pannonian Basin. The section studied is located near the Apuseni Mountains (Romania). It can be dated by the presence of foraminifera to the Varidentella reussi and Elphidium reginum biozones. The fish fauna is thus of late Serravallian/earl...

The Cenozoic growth of the intra-continental Tian Shan Range initiated during the late Eocene–Oligocene, and led to a tectonic reactivation of the complex Palaeozoic and Mesozoic lithospheric structure. Due to the very low erosion rates linked to the semi-arid climate that characterised the Tian Shan region during most of the Cenozoic, the topograp...

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)....

Among the species of Aphanius Nardo, 1827, Aphanius dispar (Rüppell, 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...

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 except...

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...

We obtained (U–Th)/He formation ages and cosmogenic ³He concentrations for pisoliths from a paleosol of the Bohnerz Formation (Siderolithic) of Central Europe. The paleosol is exposed in the Almenbühl quarry near Lohn, Canton Schaffhausen, Switzerland. The paleosol consists of red clay of 3–4 m thickness developed on deeply weathered Jurassic limes...

The highly diverse tropical freshwater fish family Cichlidae is sparsely represented in the fossil record. Here we describe the new cichlid †Tugenchromis pickfordi, gen. et sp. nov., from the Upper Miocene (9–10 Ma) of central Kenya. The new taxon possesses a unique combination of characters, including six lateral line foramina on the lacrimal, thr...

Accurate reconstruction of the final sea retreat from the North Alpine Foreland Basin (NAFB) during the Burdigalian (Early Miocene) is hampered by a lack of reliable age constraints. In this high resolution magnetostratigraphic study we try to solve a significant age bias for the onset of the Upper Freshwater Molasse (OSM) deposition in the neighbo...

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...

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 in...

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...

Here, we present the continental aquatic and terrestrial gastropods found in samples of 11 new boreholes in the Molasse Basin, southern Germany. The samples come from the Lower Freshwater Molasse (USM), the Upper Brackish Molasse (OBM; Grimmelfingen and Kirchberg Formations) and the Upper Freshwater Molasse (OSM). The studied segments of these lith...

Presented is a phylogenetic reconstruction of mudskippers using sequences of the mitochondrial COI and cytb genes. The phylogeny combines the sequences of 12 species from GenBank with those of three species (Boleophthalmus dussumieri, Periophthalmus waltoni and Scartelaos tenuis) collected from the coastal area of the Persian Gulf and Oman (Makran)...

The Miocene epoch was a time of major change in the East African Rift System (EARS) as forest habitats were transformed into grasslands and hominids appeared in the landscape. Here we provide new sedimentological and palynological data on the middle–upper Miocene Ngorora Formation (Tugen Hills, Central Kenya Rift, EARS), together with clay mineral...

Gobiiformes are one of the most versatile groups among teleosts and show countless adaptations. Their fossil record encompasses numerous otolith-based species, but only a few species that are based on skeletons, and fossil skeletons with otoliths preserved in situ are exceptionally rare. Here, articulated skeletons, some with otoliths in situ, from...

To date, 28 species of Aphanius have been recognized in the World, of which 14 species are described from six endorheic and eight exorheic basins in Iran. Here we report the taxonomic status and phylogenetic relationships of the 14 Iranian Aphanius species to understand their evolutionary history, and thus to contribute to future conservation strat...

Identification of fossil cichlids is difficult, because the currently used diagnostic morphological characters for living cichlids are mostly soft tissue based and such characters are hardly preserved in fossils. During our recent fieldwork in the Central Kenya Rift (E-Africa), we discovered several exceptionally well-preserved fossil cichlids, whi...

African freshwater cichlids (Cichlidae: Pseudocrenilabrinae) are well known for their exceptionally great diversity and their capability of rapid speciation as well as diverse adaptations. The extant Pseudocrenilabrinae can be grouped into 27 tribes, with more than 2000 species harbored in the Great Lakes and surrounding water bodies of the East Af...

Otoliths are small structures in the skull of fishes. They are responsible for hearing and orientation in the 3-dimensional space. They also hold valuable information regarding the taxonomy. Their outline, the shape of the sulcus and other features allow the determination of a fish even to the species level. A lot of fossil species are solely based...

Members of the aplocheiloid killifish genus Nothobranchius are small (3–15 cm), have a short lifecycle (3–12 months) and are widely distributed in tropical and subtropical Eastern and Central Africa. About 62 valid species of Nothobranchius are currently known. Their diversification is apparently exclusively allopatric, but up to four species of No...