Miocene Locality Research Articles

The first lacertid and partly articulated snake from the Middle Miocene of the Devínska Kobyla Hill in Slovakia, from the type locality of the earliest known seal

I here describe lacertid and snake remains from the classic Middle Miocene (MN 6) locality Bonanza of the Devínska Kobyla Hill near Bratislava (Slovakia). The locality is famous for being a type locality of an early seal Devinophoca Koretsky & Holec, 2002. During the Middle Miocene, the area of Devínska Nová Ves was part of an archipelago in the western part of the Central Paratethys in the northern part of the Vienna basin. The fossils described here comprise an incomplete left maxilla of a lacertid lizard and an articulated portion of a vertebral column of a small colubrid snake with several ribs still attached. The maxilla is tentativelly allocated here to Lacerta Linnaeus, 1758 and is a rare instance of the occurrence of the clade Lacertidae Oppel, 1811 in this environment during the Middle Miocene. Moreover, it represents the oldest known occurence of this clade in Slovakia. Articulated snake specimens from the Cenozoic are rare and although only party articulated, the specimen from Bonanza is therefore exceptional. On the basis of vertebrae alone, determination of fossil colubrids is very difficult. The morphology of the Bonanza specimen is most similar to the extant Dolichophis Gistel, 1868, as also to other small fossil snakes assigned to ‘Coluber’ Linnaeus, 1758 when that genus was a catch-all grade taxon recognised from both Europe and North America, notably ‘Coluber’ dolnicensis Szyndlar, 1987 and ‘Coluber’ pouchetii (Rochebrune, 1880). Because the potentially closest living relative of the European Miocene ‘Coluber’ is Dolichophis, I assigned tentatively the material from Bonanza to cf. Dolichophis, rather than to the wastebasket taxon ‘Coluber’.

The easternmost occurrence of the Late Miocene schizotheriine chalicothere Ancylotherium pentelicum at the classical locality of Maragheh (Iran)

The renowned Late Miocene locality of Maragheh (northwest Iran) has yielded ample material of chalicotheres, most of which has never been described in detail. The present study concerns the taxonomic evaluation of this material and its attribution to the large schizotheriine Ancylotherium pentelicum. This allows a detailed comparison to schizotheriine material from other localities, and based on this, we provide an overview of the diagnostic features of Ancylotherium that separate it from all other chalicotheres. This improves our understanding of the type species A. pentelicum. Additionally, we discuss the palaeobiogeography of the species, which is best known from the Balkan Peninsula. Maragheh marks the easternmost occurrence of A. pentelicum, but its presence has even been suggested in Africa. Lastly, we discuss the existence of sexual size dimorphism in the species, based on the herein described material along with previously described specimens of A. pentelicum from other fossil sites, such as the famous Late Miocene localities Pikermi and Samos (Greece). Specifically, it is shown that many postcranial elements exhibit great size variability that is most probably associated with significant sexual dimorphism.

The primitive giant panda Kretzoiarctos beatrix (Carnivora, Ursidae) from the hominid locality of Hammerschmiede: dietary implications

AbstractExtant giant pandas are among the most herbivorous forms of the order Carnivora, feeding mainly on hard plant material. The first steps of their evolutionary lineage are of particular interest for our understanding of the factors that led to this specialized niche. The present work deals with newly discovered dental material of the primitive ailuropodine bear Kretzoiarctos beatrix from the Late Miocene locality of Hammerschmiede (Germany). This is the first report of the genus Kretzoiarctos outside the Iberian Peninsula, expanding its spatial range to Central Europe. All of the currently known localities with K. beatrix, from both Spain and Germany, are very similar in age (c. 11.9–11.4 Ma). The present material has distinct features that enable its taxonomic discrimination from other Miocene ursids of Europe, such as Ballusia, Ursavus, Miomaci, Agriarctos and Indarctos. A thorough comparison is conducted of all of these forms. The new specimens were used in a dental microwear texture analysis in combination with ecomorphological comparisons, to investigate the dietary habits of this primitive ailuropodine. The results suggest that Kretzoiarctos was not an eater of tough plant material like the extant giant panda but was more similar to the extant Tremarctos, exhibiting opportunistic behaviour with occasional consumption of meat.

The rise and fall of the Iberian cobras (Elapidae, Naja) in the context of their European and global fossil record

AbstractVery few remains of elapid snakes are known from the Iberian Peninsula, but these include a probable endemic extinct species of cobra, Naja iberica from the Late Miocene. We here describe isolated cobra vertebrae from several Middle–Late Miocene localities in the Vallès‐Penedès Basin (Catalonia, Spain). All of these fossils are herein referred to an indeterminate species of the genus Naja. These remains are the first conclusive evidence that cobras were present in Iberia before the Messinian Salinity Crisis, and that they persisted there throughout the Miocene (and ultimately until the Pliocene). Recently, a phylogeny of extinct Naja based on cranial and vertebral morphology recognized N. iberica as a distinct lineage separate from the Central European one, represented mainly by Naja romani. However, due to taxonomic uncertainties, it is still unclear whether Iberian cobras were all part of a single lineage or whether several Naja lineages inhabited the Iberian Peninsula. They went extinct in Iberia before the end of the Pliocene. In the Vallès‐Penedès Basin, cobras were living in a mosaic environment, surviving through different phases characterized by different environmental features.

New proboscidean specimens from the Late Miocene of Romania: the huge-sized deinothere Deinotherium proavum, the rare “Mammut” cf. obliquelophus and the first description of the shovel-tusker Konobelodon from the country

In this article, we describe so far unpublished proboscidean specimens from several Late Miocene localities of Romania. A partial mandible and the complete upper/lower cheek tooth rows of a deinothere individual from the site of Gherghești 1 belong to Deinotherium proavum and comprise one of the few examples of entire cheek tooth rows of the same individual of this species. Gherghești 1 is geographically close to Mânzaţi from where the celebrated skeleton of “Deinotherium gigantissimum” was discovered at the end of the nineteenth century, and thus further highlights the importance of Romania in the study of this emblematic deinothere. Deinotherium proavum represents the last deinothere species in Europe and corresponds to the terminal stage of the size increase characterizing the evolution of European deinotheres. Two zygodont molars are attributed to the rare “Mammut” cf. obliquelophus and add to the scarce record of “Mammut” in the Miocene of Eurasia. They document the secure presence of “Mammut” in the Miocene of Romania. The small size of the studied molars compared to known specimens of the Pliocene “Mammut” borsoni and the weak development of the distal cingulum in the lower third molars may have taxonomic and biostratigraphic importance. Furthermore, the presence of an amebelodontid is documented by a large-sized and dorsoventrally flattened lower tusk fragment that shows tubular dentine in its inner part and is attributed to the tetralophodont shovel-tusker Konobelodon. This specimen marks the first record of the genus in Romania. Finally, the biostratigraphic distribution of the taxa is discussed.

New occurrences of mammals from McKay Reservoir (Hemphillian, Oregon)

AbstractEncompassing global cooling, the spread of grasslands, and biogeographic interchanges, the Hemphillian North American Land Mammal Age is an important interval for understanding the factors driving ecological and evolutionary change through time. McKay Reservoir near Pendleton, Oregon is a natural laboratory for analyses of these factors. It is remarkable for its small vertebrate fauna including rodents, bats, turtles, and lagomorphs, but also for its larger mammal fossils like camelids, rhinocerotids, canids, and felids. Despite the importance of the site, few revisions to its faunal list have been published since its original description. We expand on this description by identifying taxa not previously known from McKay Reservoir based on specimens collected during fieldwork and through reidentification of previously collected fossils. Newly identified taxa include the borophagine canid Borophagus secundus (Matthew and Cook, 1909), the camelids Megatylopus Matthew and Cook, 1909 and Pleiolama Webb and Meachen, 2004, a dromomerycid, and the equids Cormohipparion Skinner and MacFadden, 1977 and Pseudhipparion Ameghino, 1904. Specimens previously assigned to Neohipparion Gidley, 1903 and Hipparion de Christol, 1832 lack the features necessary to diagnose these genera, which are therefore removed from the site's faunal list. The presence of Borophagus secundus, Cormohipparion, and Pseudhipparion is especially important, because each occurrence represents a major geographic range extension. This refined understanding of the fauna lays the foundation for future studies of taphonomy, taxonomy, functional morphology, and paleoecology—potentially at the population, community, or ecosystem levels—at this paleobiologically significant Miocene locality.

Chilotherium schlosseri (Weber, 1905) (Rhinocerotidae, Mammalia) from the late Miocene of the foreland of the Eastern Carpathians in Romania

The paper provides an overview of the Chilotherium Ringström, 1924 fossils from Romania, which are recorded so far in three different Upper Miocene localities of the Eastern Carpathians Foreland: mammal localities of Reghiu, Bacău and Pogana. The fossils include a partial skull, several maxillary and mandible fragments and isolated teeth. They have been partially illustrated, but never described excepting the isolated teeth of Pogana. The previous assignments were confusing, ranging from Chilotherium schlosseri (Weber, 1905) and C. cf. sarmaticum for Reghiu specimens to an indeterminate species close to C. sarmaticum Korotkevich, 1958, C. kiliasi (Geraads & Koufos, 1990), and C. kowalevskii (Pavlow, 1913) for Pogana teeth. Based on descriptions and comparisons with other Chilotherium remains, all specimens from Romania can be referred to C. schlosseri. Chilotherium schlosseri is the most widespread species of the genus, ranging from Eastern Europe to Central Asia. The presence of this genus in Romania, and more globally in Eastern Europe, attests to a transition area between the more closed and wooded environments of Western Europe (where Chilotherium is absent) to the more open ones in Asia.

Faunal Correlation of Middle Miocene Locality of Chabbar Syedan with Some Other Localities of Chinji Formation of Siwalik Group

Faunal Correlation of Middle Miocene Locality of Chabbar Syedan with Some Other Localities of Chinji Formation of Siwalik Group

The Late Miocene hominoid Ouranopithecus macedoniensis (Bonis, Bouvrain, Geraads & Melentis, 1974): maxillary deciduous dentition and virtual reconstruction of the unerupted permanent teeth

The hominoid Ouranopithecus macedoniensis (Bonis, Bouvrain, Geraads & Melentis, 1974) is known from three Late Miocene localities in Greece. All are correlated with late Vallesian, MN10; more precisely, they are dated between 9.6 and 8.7 Ma. During the last 50 years several fossils of this hominoid have been recovered. The present article describes the first specimen with upper deciduous teeth, recovered from the locality Ravin de la Pluie in Axios Valley (Macedonia, Greece). The upper deciduous dentition of O. macedoniensis is characterised by a small canine relative to the length of the deciduous premolars, with a rounded occlusal crown outline, a trapezoidal dP3 with metacone and a rudimentary hypocone, a sub-squared dP4, and the absence of cingulum. The taxonomic and sexual attribution of the studied material is based on the preserved and virtually reconstructed permanent dentition, using high resolution microcomputed tomography. The morphological and metrical comparisons of the permanent dentition with the available sample of O. macedoniensis, as well as the monospecific character and the strong sexual dimorphism of the Ravin de la Pluie sample, allow its attribution to a female sub-adult individual of this taxon. The upper deciduous teeth of Ouranopithecus macedoniensis preserve some morphological features, that may have a phylogenetic value, but this hypothesis requires further investigation as the available material of all fossil taxa is poor.

New lizard material from two Early Miocene localities in France: Montaigu-le-Blin (MN 2) and Crémat (MN 3)

Squamate faunas from the MN 1–3 intervals (earliest Miocene) are scarcely documented from the European continent. I here describe squamate faunas from two French localities – Montaigu-le-Blin (MN 2) and Crémat (MN 3), the latter being the youngest locality of the Phosphorites du Quercy. The palaeobiodiversity of squamates from these sites is low relative to the faunas described from coeval localities of Amöneburg (MN 2) in Germany and Merkur-North (MN 3) in the Czech Republic. The beginning of the Miocene represents the temporary return to a paratropical humid climate after the relatively cool and dry Oligocene, and the Montaigu and Crémat materials provide previously undocumented components of herpetofaunas and their changes during this crucial time interval in France. The importance of fossil squamates from the area of the Phosphorites du Quercy is therefore highlighted for the beginning of the Miocene as well as the better known Eocene-Oligocene sites. The lizard material of Montaigu includes lacertids and blanids, the latter being represented by Blanus cf. gracilis, one of the oldest records of the genus. Moreover, it shows a higher spatial and temporal distribution of this amphisbaenian species during the Early Miocene than previously known. The material from Crémat consists of few elements which can be allocated to anguids and potentially to lacertids. Detailed figures of the specimens are provided through the means of both photography and micro-CT scanning.

Hyaenid remains from the Late Miocene of Kutschwan (Shanxi Province, China)

The hyaenid family reached its peak of spatial range and taxonomic diversity during the Late Miocene, including a very wide range of different forms that were spanning throughout the Old World. Particularly, the fossil record of the family in the Balkans and China has been extremely variable during that time. The present paper deals with cranial and postcranial hyaenid material from the Late Miocene locality of Kutschwan (Shanxi Province, China). The specimens were identified as Hyaenictitherium wongii, a widespread species with well-known occurrences throughout Eurasia during the Vallesian and the Turolian. The material presents some notable features, such as the enlarged M1, M2 and m2 that point towards a more basal form of the species. However, the statistical analysis of the variability of these characters points towards intraspecific divergence based on selective release, instead of taxonomically significant adaptive traits. The postcranial material conforms very well to the known material of H. wongii from other localities, indicating an agile cursorial hunter. This species is the only carnivoran found in Kutschwan, based on the current data.

The largest record of the minute beaver Euroxenomys minutus (Mammalia, Castoridae) from the early Late Miocene hominid locality Hammerschmiede (Bavaria, Southern Germany) and palaeoecological considerations

ABSTRACT In this study, we describe the dental remains of a very minute beaver from the early Late Miocene locality Hammerschmiede (MN 7/8) located at the Northern Alpine Foreland Basin (Southern Germany, Bavaria). The finds represent the so far most comprehensive (>1.000 dental specimens) collection of the trogontheriine castorid Euroxenomys minutus. Metrically and morphologically, the large data set demonstrates an extensive intraspecific variability of this beaver from Hammerschmiede that covers most finds of other European localities and confirms previous species assignments. The subspecies Euroxenomys minutus rhenanus from Dorn-Dürkheim (Germany), clearly isolates from this range of variation and can thus be validated. The performed age-frequency distributions for Euroxenomys minutus from the local stratigraphic levels HAM 5 (rivulet) and HAM 4 (river) correspond roughly and also resemble Euroxenomys from the swamp deposit Rudabánya (Hungary). Since previous mortality studies on the also occurring larger beaver Steneofiber depereti from Hammerschmiede demonstrated dominance of behavioural or internal factors, this cross-ecosystem similarity in mortality for the small beaver species is likely explained by external factors, such as predation. Supported by a taphonomic bone analysis, a multitude of predators are finally considered to prey on Euroxenomys minutus and indicate a predatory bone assemblage.

New data on the Late Miocene chondrichthyans from the Western Mediterranean region (Alcoy Basin, Eastern Spain)

ABSTRACT The Mediterranean Basin has been subjected to important geological and paleogeographic events that have shaped the evolution of the faunal assemblages that inhabit it. Two main events can be highlighted, the closure of its eastern margin by the Gomphotherium Land Bridge, and the temporal closure of the connection to the Atlantic Basin during the Messinian Salinity Crisis. These two events shaped the diversity of the chondrichthyan assemblages of the Mio–Pliocene Mediterranean, and their consequences can be observed even in the modern faunal composition. To better understand the taxonomic and ecological composition of the Mediterranean chondrichthyan faunas between the aforementioned events, and how they affected their evolutionary dynamics, we have performed a complete systematic and paleoenvironmental study of the chondrichthyan assemblage recovered from various Late Miocene localities at the municipality of Alcoy Basin (Alicante, eastern Spain). Our analysis allowed us to recognize up to 20 taxa of sharks and rays recovered from Miocene deposits. In addition, we reconstruct the paleobathymetry of the El Muro fossil site, based on the estimation of the Oceanization Index for foraminifera, as well as by the elasmobranch fossil assemblage via weighted bootstrap analysis. We expect that this first description and study of the Miocene chondrichthyan taxa from this area of eastern Spain could help us create a comparative framework to compare these communities with other areas, as well as show the changes in the elasmobranch species that have inhabited the Mediterranean Basin during and after the Messinian Salinity Crisis.

The late Miocene Erinaceidae and Dimylidae (Eulipotyphla, Mammalia) from the Pannonian region, Slovakia

Abstract The families Erinaceidae and Dimylidae are represented in the late Miocene localities of Slovakia (Borský Svätý Jur, Krásno, Pezinok, Šalgovce, Studienka, and Triblavina) by at least six hedgehog species—‘Schizogalerix’ voesendorfensis (Rabeder, 1973); Schizogalerix cf. S. moedlingensis (Rabeder, 1973); Lantanotherium sanmigueli Villalta and Crusafont, 1944; Atelerix cf. A. depereti Mein and Ginsburg, 2002; Atelerix aff. A. depereti, cf. Postpalerinaceus sp. indet., and Erinaceinae gen. indet. sp. indet.—and two dimylid species—Plesiodimylus chantrei Gaillard, 1897; and Metacordylodon aff. M. schlosseri (Andreae, 1904). Material of L. sanmigueli from the western Carpathians was investigated, revealing broad variability in all samples. Additionally, the deciduous premolars of Lantanotherium Filhol, 1888 are described here for the first time. Erinaceid species are frequent in the Vallesian but their abundance strongly declined afterward. As an exception, Schizogalerix Engesser, 1980 re-entered the Danube and Vienna basins during MN11, likely from eastern Europe. Members of Erinaceinae display low diversity during the late Miocene of central Europe, which tends to support a pan-European diversity phenomenon. The humidity-dependent Dimylidae spp. were abundant during the late Vallesian. Rare finds of Plesiodimylus Gaillard, 1897 confirm the survival of this family into the early MN11 in the Pannonian region.

New fossil Cetaceans from the Middle Miocene of Cuba

Remains of cetaceans, particularly odontocetes, are scarce in the Cuban fossil record. Here we describe the second odontocete (toothed whale) record from the Neogene of Cuba and extends the temporal distribution of this group in the Insular Caribbean into the Middle Miocene. This account is based on two isolated teeth found in limestones of the Güines Formation that crop out in a quarry near the town of Guanábana, Matanzas City, northwestern Cuba. The specimens are referred to the superfamily Physeteroidea incertae sedis. These findings, along with other fossils recovered from other Early and Middle Miocene localities in the region support the hypothesis of a complex, diverse marine vertebrate fauna on the formerly called “Habana-Matanzas channel” and circum-Cuban waters during the Neogene.

Unraveling the diversity of early felines: a new genus of Felinae (Carnivora, Felidae) from the Middle Miocene of Madrid (Spain)

ABSTRACT We describe a hemimandible of a medium-sized feline from the Middle Miocene (middle Aragonian, MN 5, local biozone Dc, around 15.5 Ma) site of Príncipe Pío-2, a recently discovered fossil locality placed in the urban limits of Madrid city (Spain), within an area previously known for its rich paleontological findings. The specimen is very well preserved, showing the complete hemimandible, as well as all the teeth except the incisors. Some characters, such as the missing talonid in m1, the high p3, the large mesial cuspid in p4, and the presence of a vestigial m2, prevent its adscription to other known feline taxa such as Styriofelis turnauensis or Miopanthera lorteti, previously recorded in the Middle Miocene faunas from Europe, but absent in the central basins of Spain. The hemimandible has robust mandibular corpus and ramus compared with these latter and to extant felines of similar size, and exhibits a relatively stronger ridged masseteric line, which points towards the existence of a proportionally larger and more powerful m. masseter that generated a strong bite when hunting. These characters suggest that the Príncipe Pío-2 feline could have preyed upon relatively larger prey than those of extant, similarly sized felines. We propose the new genus and species Magerifelis peignei for the studied specimen, a species that would also include the material from the French Middle Miocene localities of Bézian and Artenay.

New evidence for the unique coexistence of two subfamilies of clawed perissodactyls (Mammalia, Chalicotheriidae) in the Upper Miocene of Romania and the Eastern Mediterranean

Chalicotheres are a peculiar group of large herbivorous mammals, closely related to extant tapirs, rhinoceroses, and horses, but with large claws instead of hooves. The family Chalicotheriidae consists of two subfamilies, the Schizotheriinae and the Chalicotheriinae. Herein we present chalicothere remains from the Upper Miocene locality of Pogana 1 in Romania, identifying the schizotheriine Ancylotherium pentelicum and an indeterminate chalicotheriine that were both found in the same stratigraphic layer. Thus, the Pogana 1 locality represents one of the very few confirmed cases of the co-occurrence of the two subfamilies within one fossiliferous horizon in the same fossil site. A detailed review of all localities where the two subfamilies have been reported to co-occur shows that this is a rare phenomenon that is almost exclusively observed in the Turolian of the Balkan-Iranian zoogeographical province. This is probably due to provincial differences in the palaeoenvironment. The data presented here support the hypothesis of a diverse mosaic environment in the Balkan-Iranian province with both closed environments and open habitats that was able to sustain a rich and diverse large mammal fauna.

An enigmatic giraffid from the latest Miocene of Italy: Taxonomy, affinity, and paleobiogeographic implications

The origin and evolution of endemic species characterizing the Oreopithecus-faunal assemblages of the Tusco-Sardinian archipelago remain a matter of debate. An emblematic case is the enigmatic giraffid Umbrotherium azzarolii, represented by a single specimen from the type locality of Casteani (Tuscany) and by several isolated teeth and fragmentary mandibles from the locality of Fiume Santo (Sardinia). An exhaustive diagnosis of Umbrotherium has not been firmly established, and its systematic and phylogenetic position remain unresolved. Unpublished remains of giraffids, including an almost complete mandible, several isolated teeth, and other cranial remains are described for the first time in the present work. The specimens were collected from the locality of Botro della Canonica (Pisa), located at the northernmost portion of the Tusco-Sardinian archipelago. The new material sheds light on the morphological and morphometric variability of Umbrotherium, thereby enabling a comparison between specimens collected from different Tusco-Sardinian Miocene localities spanning from the V1 to the V2 Oreopithecus-Zone Faunas and allowing the establishment of the new species U. engesserii sp. nov. from Fiume Santo (Sardinia). This study also reveals that Umbrotherium was more closely related to Decennatherium than to other Late Miocene continental giraffids, suggesting a dispersal of its ancestor from the Iberian Peninsula. Accordingly, a new paleogeographic and biochronological framework is proposed herein for the Tusco-Sardinian archipelago, hypothesizing a fragmentation of the area into several domains, with sporadic reconnections, and the establishment of different faunal assemblages.

Middle Miocene “Giraffa” from the Chinji Formation of Siwaliks, Pakistan

A comprehensive record of unique and endemic species of “Giraffa” priscilla has been documented and reported from the Chinji Formation of Lower Siwaliks, Pakistan. The material comprises incisors, canines, premolars, molars, mandibular and maxillary fragments which have been recovered from the Middle Miocene localities of Dhok Bun Ameer Khatoon, Rakh Wasnal, Bhelomar, and Lawa, Punjab, Pakistan. “Giraffa” priscilla is a primitive species of giraffids, having a broad crown, strong styles, and median ribs.

First Mexican records of Anthracotheriidae (Mammalia: Artiodactyla)

ABSTRACTAnthracotheres are generalised artiodactyls that have an extensive record in the Cenozoic of Eurasia and Africa. In North America they have been collected in middle Eocene to early Miocene localities from the California Coast, the Great Plains and the Gulf Coast of the United States, with a single record from the early Miocene of Panama. Here we report few specimens from the early Oligocene (Ar1) Iniyoo Local Fauna of north-western Oaxaca, and the earliest Miocene of Simojovel de Allende, in northern Chiapas. This material has diverse features that indicate they belonged to the bothriodontineArretotherium, such as selenodont cristids associated with the protoconid and hypoconid, the absence of a premetacristid, and the crenulated enamel. They share withArretotherium acridensandArretotherium meridionalethe absence of a mesiolingual metacristid, but their general morphology and size indicate a close relationship toAr. meridionale. Nevertheless, in absence of better-preserved specimens, we decided not to assign the fossil material to this species. Specimens from Oaxaca and Chiapas are the first records of anthracotheres in Mexico. These new records link the previous ones from temperate North America and tropical Central America and indicate that Anthracotheriidae had a very wide geographical distribution in North America during the Palaeogene and the Neogene. Additionally, they represent the southern-most records ofArretotheriumin North America during the Oligocene and the early Miocene.