Phylogeographical and speciation patterns in subterranean worm lizards of the genus Blanus (Amphisbaenia: Blanidae)

The peculiar lifestyle of subterranean reptiles must determine their modes of speciation and diversification. To further understand the evolutionary biology of subterranean reptiles, we studied the phylogeny of worm lizards of the genus Blanus and the phylogeography of its Iberian representatives. We used mitochondrial (ND4 and 16S rRNA) and nuclear (anonymous) partial gene sequences to resolve phylogenetic relationships within Blanus. The Eastern Mediterranean Blanus strauchi was recovered as sister group of Western Mediterranean species. Iberian and North African Blanus were recovered as reciprocally monophyletic groups. The same genes were used to determine phylogeography of 47 populations of Blanus cinereus. Mitochondrial and nuclear sequence data recovered two highly supported Iberian clades. Parapatry and high sequence divergences between them suggest that these clades may represent independent taxonomic units. A molecular clock was calibrated considering that the split between Iberian and North African Blanus was due to the re-opening of the Betic Strait in the Upper Tortonian (8-9 million years ago). Differentiation between the two Iberian clades was estimated to date back to 5.2 million years ago. The Central Iberian clade included five mitochondrial haplotype lineages (A-E). Geographical ranges of two of them broadly overlap in the central Iberian plateau. After testing alternative hypotheses, the most likely explanation for this striking phylogeographical pattern involves recent dispersal of one of the lineages (C) over the geographical range of the other (B). The inferred recent dispersal of this fossorial reptile is explained in terms of demographic advantages associated to underground lifestyle.     

The species identity and biogeography of Blanus (Amphisbaenia: Blanidae) in Lebanon

The genus Blanus Wagler, 1830 represents limbless, burrowing reptiles of the family Blanidae with disjunct circum-Mediterranean distribution. The recently described species Blanus alexandri Sindaco, Kornilios, Sacchi & Lymberakis, 2014 is known from south-eastern Turkey with a presumed occurrence in the Levant and Iraq. We provide here records from Lebanon and confirm the affiliation of Lebanese populations to this species by mitochondrial and nuclear DNA. Blanus alexandri comprises at least seven deeply evolved phylogenetic clades with up to 11% of uncorrected p-distances in their mitochondrial DNA. This suggests a probably older than the Miocene origin of some of these clades. Populations from Lebanon form a different clade with a genetic diversity that is close to populations from southern Turkey.     

A contribution to the herpetofauna from the late Miocene of Gritsev (Ukraine)

Excavations in the locality Gritsev (karstic fillings of the upper Miocene age in a limestone quarry in Shepetovsky District, Khmelnitsky Region, Ukraine) revealed, since 1983 when excavations started, a rich vertebrate fauna that includes also amphibians and lizards. The material is rich enough to assess individual and developmental variation within some taxa (e.g., Pseudopus pannonicus). Although the fragmentary nature of the disarticulated bones does not sometimes allow precise taxonomic assignment, morphological differences in, e.g., dental characteristics suggest the presence of some other ones, not formally named forms. Thus, the list of the amphibian and reptilian taxa from Gritsev is by far not complete, but their osteological characteristics are representative to such a degree that they can provide information for comparisons with other European Miocene herpetofaunas.     

Orbital signatures in a late Miocene dinoflagellate record from Crete (Greece)

A high-resolution palynological study of the cyclically bedded Faneromeni section (upper Tortonian-lower Messinian) on Crete (Greece) is presented. This study aims to recognize orbitally-driven variations in the palynological record and to validate the age model based on the astronomical calibration of the sedimentary cycles. Four palynology-based environmental proxies were utilised using interpretations of fossil dinoflagellate associations based on modern ecological characteristics. Cross-spectral analysis between the proxy records and astronomical target curve, the 65°N summer insolation, yielded in most cases significant spectral power and coherence in the precession and/or obliquity frequency bands. Precession-controlled variations in the proxy records are related to lithology and indicate that maxima in continental input and minima in sea surface salinity coincide with sapropel formation. The influence of obliquity is most clearly reflected in the index of continental versus marine palynomorphs (S-D). The absence of a distinct time lag relative to obliquity indicates that the 41-kyr component in continental input is controlled by oscillations in regional Mediterranean climate rather than by glacial cyclicity. Phase relations in the different astronomical frequency bands of the spectrum, as compared with the Mediterranean Pliocene, essentially confirm the validity of the Miocene astronomical time scale. Finally, a major non-cyclic change in the palynological assemblage at 6.68 Ma indicates enhanced salinity and decreased river discharge. This shift coincides with a significant drop in sedimentation rate informally termed the “Early Messinian Sediment starvation Event”.     

Muridae from the late Miocene site of Venta del Moro (Eastern Spain)

Murids are usually the dominant faunas of the late Miocene and early Pliocene micromammal assemblages. The present work deals with the murid faunas of the well-known late Miocene locality of Venta del Moro, comprising over 2700 molars ascribed to the taxa Apodemus gorafensis, Occitanomys alcalai, Paraethomys meini and Stephanomys dubari. This list differs from those of previous studies by the presence of A. gorafensis instead of A. gudrunae. The presence of A. gudrunae in younger localities than Venta del Moro implies that both species, considered ancestor and descendant, coexisted for some time. In addition, the analysis of such an extensive collection has allowed us to check the variability of the studied species in great detail, such as the development of the longitudinal connections of O. alcalai, to a greater extent than previously described. Finally, we discuss the biometrical changes of the lineages of Paraethomys in Southern Europe from MN13 to MN15, and hypothesize about the mechanisms behind these variations in size.     

The first known fossil record of pygmy pipehorses (Teleostei: Syngnathidae: Hippocampinae) from the Miocene Coprolitic Horizon,Tunjice Hills,Slovenia

The first known fossil record of pygmy pipehorses is described. The fossils were collected in the Middle Miocene (Sarmatian) beds of the Coprolitic Horizon in the Tunjice Hills, Slovenia. They belong to a new genus and species Hippotropiscis frenki, which was similar to the extant representatives of Acentronura, Amphelikturus, Idiotropiscis, and Kyonemichthys genera. Hippotropiscis frenki lived among seagrasses and macroalgae and probably also on a mud and silt bottom in the temperate shallow coastal waters of the western part of the Central Paratethys Sea. The high coronet on the head, the ridge system and the high angle at which the head is angled ventrad indicate that Hippotropiscis is most related to Idiotropiscis and Hippocampus (seahorses) and probably separated from the main seahorse lineage later than Idiotropiscis.     

Anatomical study of the skull of amphisbaenian Diplometopon zarudnyi (Squamata,Amphisbaenia)

This study investigates the amphisbaenian species skull which includes cranium, lower jaw and hyoid apparatus. The medial dorsal bones comprise the premaxilla, nasal, frontal and parietal. The premaxilla carries a large medial tooth and two lateral ones. The nasals are paired bones and separated by longitudinal suture. Bones of circumorbital series are frontal, orbitosphenoid and maxilla. The occipital ring consists of basioccipital, supraoccipital and exooccipital. Supraoccipital and basioccipital are single bones while the exo-occipitals are paired. The bones of the palate comprise premaxilla, maxilla, septomaxilla, palatine, pterygoid, ectopterygoid, basisphenoid, parasphenoid, orbitosphenoid and laterosphenoid. Prevomer and pterygoid teeth are absent. Palatine represent by two separate bones. The temporal bones are clearly visible. The lower jaw consists of the dentary, articular, coronoid, supra-angular, angular and splenial. The hyoid apparatus is represented by a Y-shaped structure. The mandible is long and is suspended from the braincase via relatively short quadrate. There is an extensive contact between the long angular and the large triangular coronoid. Thus inter-mandibular joint is bridged completely by the angular and consequently, the lower jaws are relatively rigid and kinetic. The maxillae are suspended from the braincase largely by ligaments and muscles rather than through bony articulation. In conclusion, the skull shape affects feeding strategy in Diplometopon zarudnyi. The prey is ingested and transported via a rapid maxillary raking mechanism.     

A fossil Vallisneria plant (Hydrocharitaceae) from the Early Miocene freshwater deposits of the Most Basin (North Bohemia)

A new fossil species of Vallisneria (V. janecekii Bogner & Kva?ek, sp. n., Hydrocharitaceae) is reported from Early Miocene freshwater lake deposits of the North Bohemian brown coal basin (Czech Republic). It is based on a single fragmentary sterile plant bearing two clusters of ribbon-shaped widely serrulate leaves on a short rhizome. It is a potential producer of seeds described from equivalent and slightly older deposits in Saxony (Germany) as Vallisneria ovalis Mai.     

A Miocene leaf fossil record of Rosa (R. fortuita n. sp.) from its modern diversity center in SW China

Roses (Rosa, Rosaceae) are arguably the most admired ornamental plants in the world. Southwestern China is the center of diversity for many extant native species of Rosa and fossils found in this region are critical for understanding the evolution of this genus. Herein, we report a leaf fossil record with good preservation from the late Miocene of Yunnan Province. The opposite and odd-pinnate leaf is composed of seven elliptical leaflets, with close, crenulate, and regular marginal teeth. The stipules are lanceolate and adnate to the petiole. Additionally, the secondary veins are semicraspedodromous, showing the same venation pattern as most living Rosa species in southwestern China. On the basis of the extensive morphological comparisons, we propose a new species, R. fortuita T. Su et Z.K. Zhou n. sp. This is the first confirmed Rosa leaf fossil record in China, and the discovery of R. fortuita n. sp. indicates that Rosa existed in southwestern China by the late Miocene. It suggests that Rosa was distributed in subtropical or temperate forests and shared a similar ecological niche as Rosa in Europe during the Oligocene and Miocene. The modern diversification of Rosa in southwestern China is thought to have been closely associated with the continuous uplift of the Qinghai-Tibet Plateau since the late Miocene, creating complex topography and a variety of climate conditions.     

A new fossil Aphanius species from the Upper Miocene of Armenia (Eastern Paratethys)

A new fossil fish species, Aphanius yerevanicus (Cyprinodontiformes, Cyprinodontidae), is described based on a well-preserved, articulated skeleton from the early Upper Miocene of Yerevan, Armenia (Eastern Paratethys). The fish-bearing sediments contain a fossil assemblage indicative of a euryhaline environment. The new species differs from previously described fossil Aphanius species by the possession of caudal ribs articulating with the first haemal spine, a remarkably low number of abdominal vertebrae, and the morphology of the tricuspid jaw teeth. Co-occurrence of A. yerevanicus with Prolebias mutilus Bogachev, 1936 at Yerevan suggests that sympatric occurrence of several members of the Cyprinodontiformes continued successfully throughout the Miocene in euryhaline environments where competition between these forms and other fishes was low because of the unstable salinity.     

The first known fossil record of pipehorses (Teleostei: Syngnathidae: Haliichthyinae) from the Miocene Coprolitic Horizon from the Tunjice Hills,Slovenia

The first known fossil specimens of pipehorses (Haliichthyinae) were unearthed from the Middle Miocene (Sarmatian) beds of the Coprolitic Horizon in the Tunjice Hills, Slovenia. These fossil pipehorses belong to a new genus and species Hippohaliichthys edis, which was similar to the extant species Haliichthys taeniophorus. The body morphology indicates that the described fossil pipehorses were also closely related to the pygmy pipehorse Hippotropiscis frenki and the seahorse Hippocampus sarmaticus, two taxa which were also found in the Coprolitic Horizon. The described fossil material of pipehorses indicates that seahorses evolved from a group of pipehorses that were similar in size and shape to extant and fossil pipehorses of the Haliichthyinae subfamily.     

Dentition ofSivaladapis nagrii (Adapidae) from the late Miocene of India

Two genera and three species of adapid primates are known from the middle and late Miocene of India and Pakistan. Most fossil specimens are fragmentary, but the best-known species, Sivaladapis nagrii,is now represented by enough specimens to permit composite reconstruction of much of the dentition. The incisors of Sivaladapishave spatulate crowns, and the canines are large, projecting teeth. Premolars and molars exhibit complex occlusion involving simultaneous approximation of pointed leading cusps on upper and lower molars, with linear trailing lophs. The premolar eruption sequence in Sivaladapisappears to be P ₂-P₄-P₃, as in most extant prosimians. Symphyseal fusion of the mandibular rami occurred early in ontogeny, before the eruption of any of the anterior permanent teeth. We interpret Sivaladapisto have been a specialized arboreal folivore that became extinct near the end of the Miocene, when the distribution of forests was increasingly restricted and colobine monkeys first invaded South Asia.     

Brachypotherium (Perissodactyla,Rhinocerotidae) from the late Miocene of Samburu Hills,Kenya

Several isolated cheek teeth and mandibular specimens of Rhinocerotidae (Mammalia, Perissodactyla) from the upper Miocene Namurungule Formation in Samburu Hills, Kenya, are redescribed. Previously, these specimens had been identified as Chilotheridium pattersoni, Chilotheridium sp., Paradiceros mukirii, and Paradiceros sp. They are reidentified here as documenting the genus Brachypotherium based on their bucco-lingually broad molariform upper premolars with short crochet and flattened buccal walls on both upper and lower molars, the latter having a shallow external groove. Comparisons with other Brachypotherium species suggest that the present specimens belong to Brachypotherium sp. cf. B. minor. The presence of Brachypotherium in the Samburu Hills, at ca. 9.5 Ma, is concordant with the paleoenvironment (presence of lacustrine and river environments) known for this locality during the early late Miocene.     

Miocene Ethiopian amber: A new source of fossil cryptogams

Amber is renowned for the exceptional preservation state of its inclusions, allowing detailed morphological analysis and providing relevant environmental, palaeoecological, geographical, and geological information. Amber deposits are predominantly known from North America, Europe, and Asia, and are considered to be rare on the continents that formed Gondwana. The recent discovery of fossiliferous amber deposits in Ethiopia, therefore, provides an inimitable opportunity to close gaps in the fossil record of African terrestrial biota and to study organisms which are otherwise rare in the fossil record. Here we show that diverse cryptogams are preserved in highest fidelity in Miocene Ethiopian amber. We describe gametophyte fragments of four liverworts: Thysananthus aethiopicus sp. nov. (Porellales, Lejeuneaceae), Lejeunea abyssinicoides sp. nov. (Porellales, Lejeuneaceae), Frullania shewanensis sp. nov. (Porellales, Frullaniaceae), and Frullania palaeoafricana sp. nov. (Porellales, Frullaniaceae). Furthermore, we describe a pleurocarpous moss of the extant genus Isopterygium (Hypnales, Pylaisiadelphaceae) and a lichen representing the order Lecanorales. These new specimens represent the first amber fossils of liverworts, mosses, and lichens from the African continent and render Ethiopian amber as one of the few worldwide amber deposits preserving bryophytes (mosses and liverworts) or lichens. Fossil species of Thysananthus were recorded in Eocene Baltic and Oligocene Bitterfeld as well as Miocene Dominican and probably also Miocene Mexican ambers. Fossils that can unequivocally be assigned to Lejeunea have only been found in Dominican amber so far. Neotropical ambers contain only one taxon of Frullania to date, while the genus is most diverse in Baltic, Bitterfeld, and Rovno ambers, formed in temperate regions. The new fossils support a tropical to subtropical origin of Ethiopian amber. The new African liverwort fossils are included in an updated list of leafy liverworts described from worldwide Cenozoic ambers to date.     

A diverse fossil terrestrial arthropod fauna from New Zealand: evidence from the early Miocene Foulden Maar fossil lagerstätte

Fossil evidence for the evolutionary history of terrestrial arthropods in New Zealand is extremely limited; only six pre‐Quaternary insects (Triassic to Eocene) have been recorded previously, none of Miocene age. The Foulden Maar fossil lagerstätte in Otago has now yielded a diverse arthropod assemblage, including members of the Araneae, Plecoptera, Isoptera, Hemiptera, Coleoptera, Hymenoptera, Trichoptera and Diptera. The fauna significantly emends the fossil record for the Southern Hemisphere, provides an unparalleled insight into a 23‐million‐year‐old New Zealand lake/forest palaeoecosystem and allows a first evaluation of arthropod diversity at a time coeval with or shortly after the maximum marine transgression of Zealandia in the late Oligocene. The well‐preserved arthropods chiefly represent ground‐dwelling taxa of forest floor and leaf litter habitats, mostly from sub‐families and genera that are still present in the modern fauna. They provide precisely dated fossil evidence for the antiquity of some of New Zealand's terrestrial arthropods and the first potential time calibrations for phylogenetic studies. The high arthropod diversity at Foulden Maar, together with a subtropical rainforest flora and fossil evidence for complex arthropod–plant interactions, suggests that terrestrial arthropods persisted during the Oligocene marine transgression of Zealandia.     

Rodents and insectivores from the late Miocene of Romerales (Fortuna Basin,Southern Spain)

Abstract

The Neogene Fortuna Basin (Murcia Region, SE Spain) is rich in microvertebrate sites. Its continental sections include localities extended from the middle Turolian (MN12) to the early Ruscinian (MN14). However, there are few works dealing with the taxonomy of these fossil assemblages. In this paper we provide a complete taxonomic study of the rodents and insectivores from three levels (ROM-2B, ROM-2C and ROM-3A) within the Romerales section. We infer a late Turolian age (late Messinian) for these levels, among which the richest and most diverse level is ROM-C, including at least 11 different taxa. In addition, the paleoecological analysis of these fossil assemblages suggests the dominance of open herbaceous meadows under temperate climate during the formation of these sites, with a slight decrease in temperature and humidity from ROM-2B to ROM-2C.     

Musteloids and feloids (Mammalia, Carnivora) from the late Miocene locality of Pikermi (Attica, Greece)

Some relatively rare carnivore remains from Pikermi are studied. The present specimens include Simocyon primigenius(ROTH and WAGNER, 1854), Promeles palaeatticus(WEITHOFER, 1888), Martes woodwardiPILGRIM, 1931, Felis atticaWAGNER, 1857 and Machairodus giganteus(WAGNER, 1848), as well as some postcranial remains that resemble Enhydriodon? latipesPILGRIM, 1931 and are assigned to Mustelidae gen. et spec. indet. Some of these species were not known until now but from their holotypes as M. woodwardi and E.? latipes. Other species such as F. attica and M. giganteus are better known but from relatively scant material. Since the material of the earlier mentioned species comes from Pikermi, their type locality, it is of especial importance and improves our knowledge for the Pikermi carnivores and their relationships to other species. In some cases where the material comes from ossiferous blocks, the accompanying faunal context is discussed.