Middle Turolian Research Articles

A newly discovered Hystrix primigenia specimen from the Kemiklitepe collection at Ege University Natural History Museum: insights into paleobiogeography in Eurasia.

Porcupines, members of the Hystricidae family, represent a unique group of herbivorous mammals. This study details the identification of a newly discovered mandible fragment of Hystrix primigenia, along with a right cheek tooth series from the middle Turolian Kemiklitepe-A fossil locality. While Hystrix fossils are found in numerous localities, the materials are often limited to a few dental fragments or isolated teeth, posing challenges to systematic investigations. The examination of this lower tooth series prompted a comprehensive review of all H. primigenia findings across Eurasia, shedding light on its adaptive characteristics over time and space. Our paleobiogeographical analysis indicates the absence of H. depereti in Late Miocene Türkiye, while the dispersal range of H. primigenia is broader than that of H. depereti in Eurasia. Additionally, the study delves into the discussion of H. primigenia and H. depereti findings in Eurasia, ultimately refining the categorization of Late Miocene Hystrix discoveries in Türkiye to two species: H. primigenia and H. kayae. Our review suggests the possibility of an additional H. kayae finding from Samos, Greece.

The Vallesian large Palaeotragus Gaudry, 1861 (Mammalia: Giraffidae) from Northern Greece

We herein re-discuss the systematics of the Late Miocene representatives of the most common but poorly documented Eurasian giraffid genus Palaeotragus on the occasion of the review and description of new samples from the Vallesian faunas of Northern Greece. Our results detect five Late Miocene Palaeotragus morphotypes, recognizing at least four species. The so called ‘small-sized palaeotrages’ are represented by the type species P. rouenii and the Chinese P. microdon, whereas the validity of P. pavlowae from Grebeniki (Ukraine) is doubted. ‘Large-sized palaeotrages’ are mainly represented by P. coelophrys (synonym of P. expectans, P. borissiaki, P. hoffstetteri, P. quadricornis, and probably P. moldavicus), a species that thrived in the peri- Black Sea territories during Vallesian and survived during Turolian in the Irano-Anatolian domain, likely by adopting a more robust appearance. Large palaeotrages from the Vallesian faunas of Pentalophos and Ravin de la Pluie (Axios Valley, Greece) are identidied as P. coelophrys with certain confidence. The Vallesian P. berislavicus from Berislav (Ukraine) has intermediate morphometric features between P. rouenii and P. coelophrys and it is, therefore, recognized as a most probably valid species. The latest Vallesian Nikiti-1 (Chalkidiki peninsula) large palaeotrage shares many morphometric features with P. berislavicus, suggesting that the species may have invaded Balkans by the end of Vallesian and possibly survived there until middle Turolian. The Late Miocene Palaeotragus asiaticus from Central Asia is a quite problematic species; it appears closely related to the Turolian equivalent P. cf. coelophrys from China and both may be linked to the older Berislav taxon.

Old world hipparion evolution, biogeography, climatology and ecology

Nearly five decades ago Berggren and Van Couvering proposed an Old World “Hipparion Datum” wherein a North American Hipparion extended its range across Eurasia and Africa as an “instantaneous prochoresis” populating the Old World. Four decades ago Woodburne and Bernor examined European and North African hipparion assemblages and proposed a number of distinct hipparion lineages, sharply departing from the mono-generic paradigm of previous work. Through the 1980s until now, hipparion systematic studies have delineated multiple superspecific groups of hipparions. Herein, we define 10 recognizable genus-rank Eurasian and African taxa delineating their chronologic occurrences, geographic extent and where data exists, their body mass and paleodietary preferences. Our study supports the current interpretation that a species of North American Cormohipparion extended its range into the Old World in the early late Miocene. Regional first occurrences of Cormohipparion are recognized in the Potwar Plateau, Pakistan and Sinap Tepe, Turkey 10.8 Ma. The slightly derived lineage Hippotherium is recorded earlier in the Pannonian C of the Vienna Basin, 11.4–11.0 Ma marking the chronologic “Hipparion” Datum at the lower boundary of Mammal Neogene (MN) Unit 9. Within MN 9, 11.2–9.9 Ma, Cormohipparion underwent a minor diversification whereas Hippotherium diversified in Central and Western Europe and China and Sivalhippus (S. nagriensis) originated in the Indian Subcontinent. Whereas Cormohipparion did not survive into the late Vallesian, MN10 (9.9–8.9 Ma), Hippotherium and Sivalhippus did and the Cremohipparion and Hipparion s.s. lineages originated. During the early and middle Turolian (MN11–12, 8.9–6.8 Ma) Hippotherium, Sivalhippus, Cremohipparion and Hipparion persisted and new lineages, Eurygnathohippus, Plesiohipparion, Baryhipparion and Shanxihippus originated. An initial extinction interval occurred at the end of the Miocene, MN13 (6.8–5.3 Ma) wherein all but one endemic species of Hippotherium, H. malpassi (Italy), Hipparion and several species of Cremohipparion became extinct. Lineage and species reduction continued across the Mio-Pliocene boundary so that by the beginning of the Pliocene (MN14, 5.3 Ma) only African species of Eurygnathohippus, Chinese Plesiohipparion houfenense and Proboscidipparion sinense remained. The later Pliocene (MN15–16, ca. 5.0–2.5 Ma) documents the persistence of endemic Chinese Baryhipparion insperatum, modest diversification of African Eurygnatohippus spp. and Chinese Plesiohipparion and Proboscidipparion spp. Eurygnathohippus made a limited geographic extension into the Indian subcontinent during MN16, whereas Pleisohipparion and Proboscidipparion extended their ranges into Eurasia during MN15 and MN16. The latest occurring hipparions are Proboscidipparion sinense at 1.0 Ma in China and Eurygnathohippus cornelianus in Africa <1.0 Ma. Old World hipparion lineages early on increased their body mass in MN9. During the Turolian interval (MN11–13) hipparion lineages diversified their body mass from very small (<100 kg) to heavy forms (>300 kg), with the smaller forms being predominately grass feeders and larger ones being mixed feeders. Decreased hipparion lineage and species diversity in the Pliocene was accompanied by increased average body size and hypsodonty probably in response to more seasonal Eurasian and African environments. There is no evidence that hipparions ever adapted to cold and dry Old World Pleistocene environments.

Large giraffids (Mammalia, Ruminantia) from the new late Miocene fossiliferous locality of Kemiklitepe-E (Western Anatolia, Turkey)

Kemiklitepe is a well-known locality with four recognised fossiliferous horizons, KTA to KTD, which have yielded a plethora of mammalian remains. Previous taxonomic studies indicate the presence of three giraffid taxa: Samotherium major and Palaeotragus rouenii from the uppermost three horizons, KTA, KTB and KTC, as well as Palaeotragus rouenii and Samotherium? sp. from the lowermost KTD horizon. In this study a new locality, Kemiklitepe-E, is presented for the first time. Kemiklitepe-E is located approximately 350 m NW of the classic Kemiklitepe locality. The fossiliferous sedimentary rocks at Kemiklitepe-E occur at the same stratigraphic level as localities KTA, KTB and KTC. The preliminary faunal list includes representatives of Proboscidea, Chalicotheriidae, Equidae, Bovidae and Giraffidae. Comprehensive descriptions and comparisons of the Kemiklitepe-E Giraffidae specimens suggest the co-occurrence of two large giraffids: Samotherium major and Helladotherium duvernoyi. Samotherium major, previously documented from this region, is the most common taxon at Kemiklitepe. Helladotherium duvernoyi is rare at Kemiklitepe and here reported for the first time. The two taxa coexisted during the middle Turolian in Greece and Western Anatolia. In addition, it is suggested that specimens of Samotherium? sp described from KTD possibly belong to Samotherium neumayri. Based on the stratigraphic position of fossiliferous rocks, as well as the faunal data presented herein, the newly discovered locality is considered to be of middle Turolian (MN12) age.

Palaeoecological and palaeoenviromental reconstruction of the upper Miocene vertebrate karstic site of Corral de Lobato, central-eastern Spain

We report stable isotopic analyses for dental enamel of fossil mammals from the Miocene (~7.1 Ma, MN13, middle Turolian) Corral de Lobato locality, Guadalajara, Spain, and compare it with adjacent localities of the Iberian Range. This multidisciplinary research allows the assessment of the palaeoenvironmental, palaeoecological and palaeoclimatic conditions that were in place in the central-eastern area of the Iberian Peninsula at this time. Herbivore tooth enamel δ13C values point to the presence of two environments: open woodland/arid C3 grassland and woodland/mesic C3 grassland. We infer the most probable diets of the two carnivores analysed, indicating a partition of resources and habitats. Tooth enamel δ18O values from the obligate drinkers indicate that all of them would have drunk water with similar evaporation rates except for the suid Hippopotamodon major, although its distinct δ18O values may be influenced by its omnivorous diet. Comparing Corral de Lobato taxa with those from other semi-contemporaneous (MN12–MN13) localities of the nearby Teruel Basin, δ13O values point to more arid conditions in Corral de Lobato while lower values of δ18O are most likely related to the karst origin of water bodies. Possibly, the difference between the isotopic compositions of Corral de Lobato and adjacent localities results from the combined effect of palaeotopography and a global climate pulse about 7 Ma.

First European Neogene record of true pheasants from Gorna Sushitsa (SW Bulgaria)

A late Miocene (Middle Turolian) ulna is described as a holotype of Phasianus bulgaricus sp. n.. This find is the first record of Neogene true pheasants in Europe.

Proboscideans from the upper Miocene localities of Thermopigi, Neokaisareia and Platania (Northern Greece)

In the present article, we study the proboscidean remains from three upper Miocene localities of Northern Greece: Thermopigi (Serres), Neokaisareia (Pieria) and Platania (Drama). The material from the Turolian locality of Thermopigi includes only postcranial specimens. The morphological features of the scapula indicate the presence of the deinotheriid Deinotherium sp., whereas the rest of the specimens are morphologically distinct from Deinotherium and can be referred to Elephantimorpha indet. The material from Neokaisareia consists of a partial skeleton of a single individual and is attributed to the mammutid Mammut sp. (M.obliquelophus?). This taxon is known in Greece from the early–middle Turolian. The Platania proboscidean belongs to the tetralophodont amebelodontid Konobelodon cf. atticus. The genus Konobelodon was already present during the Vallesian of the wider area, but the lower tusk of the Platania shovel-tusker presents some morphological and metrical differences from the Vallesian representative, yet it has also smaller dimensions in its deciduous dentition than the morphologically similar Turolian specimens. The type locality of K. atticus is Pikermi (Attica, Greece), correlated to the middle Turolian, but the known biostratigraphic range of this species covers the entire Turolian. Platania is possibly correlated close to the Vallesian/Turolian boundary and the possible record of this species could document one of its earliest occurrences.

Magnetostratigraphic dating of the late Miocene Baogeda Ula Formation and associated fauna in central Inner Mongolia, northern China

Detailed understanding to the evolution of Neogene land mammals in East Asia and its intercontinental correlation has been impeded by the absence of an integrated biochronological system of this region. The numerous and diverse records of Neogene vertebrate fossils preserved in Inner Mongolia of northern China play a key role in the establishment of an independent biochronological framework in East Asia. However, most of these faunas are poorly constrained by independent chronological controls due to the scattered distribution of fossil localities and insufficient exposure of the fossiliferous strata. Additionally, age estimates by mammalian evolution and correlation often have uncertainties greater than ~1–2 million years. Here we present new magnetostratigraphic results of the Baogeda Ula Fauna, which was generally assigned an early Baodean Age (Late Miocene), or equivalent to the European MN12 zone (middle Turolian). Hipparion remains and associated vertebrate fossils were excavated from the upper part of the fluvio-lacustrine Baogeda Ula Formation near Abaga Banner, central Inner Mongolia of northern China. At least two layers of basalt sheet flows can be observed on top of the Baogeda Ula Formation. Our magnetostratigraphy, aided by published biochronology data and K–Ar ages of the lower basalt layer suggests that the Baogeda Ula Fauna consisting of three fossiliferous horizons can be placed within chron C4n.1n with an age range of 7.642–7.528 Ma. Thus, the Baogeda Ula Fauna becomes the first and only Baodean assemblage that is constrained by both magnetostratigraphy and radiometric ages, offering an anchoring point for future biochronological correlations.

Iberian micromammals show local extent of Vallesian Crisis

Once perceived as a continent-wide extinction event, doubts have been raised as to the severity and extent of the Vallesian Crisis. In this paper, we use the high-resolution late Aragonian to middle Turolian micromammal record from both the coastal Vallès-Penedès Basin and inland Calatayud-Teruel Complex to determine geographic extent of this late Miocene turnover event and its effects within the reaches of the Iberian Peninsula. Differences in faunal composition between the two regions confirm that the event was much less severe than previously thought. Surprisingly, the diversity peaks in the two regions do not coincide. Whereas at the coast highest diversity is found in the early Vallesian (just before the turnover), in the inland it peaks in the late Vallesian. Several taxa never made it inland as reflected by the low similarity between both regions during the early Vallesian, suggesting that the Vallès-Penedès was indeed unique and consistently showed higher affinities with northern regions. The large differences even at relatively short distances support the idea that the ‘Vallesian Crisis’ was a local event.

Reprint of: The huge-sized deinothere Deinotherium proavum (Proboscidea, Mammalia) from the Late Miocene localities Pikermi and Halmyropotamos (Greece)

Presented here are the deinotheriid dental and postcranial remains from the Late Miocene localities Pikermi and Halmyropotamos (Greece). The study and comparison of the available juvenile dental material from Pikermi with other relevant specimens from Europe showed that it belongs to the huge-sized Deinotherium proavum (= D. gigantissimum). Additionally, several postcranial specimens from Pikermi, as well as from Halmyropotamos, present deinotheriid features, which distinguish them from elephantoids, and permit their attribution also to D. proavum. This species is known from the Turolian and represents the terminal evolutionary stage of the European deinotheres. Its presence in Pikermi and Halmyropotamos is compatible with the middle Turolian (MN 12) age of the localities. Additionally, the taxonomy of European deinotheres is discussed, as well as the biostratigraphical and geographical distribution of D. proavum.

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

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.

The huge-sized deinothere Deinotherium proavum (Proboscidea, Mammalia) from the Late Miocene localities Pikermi and Halmyropotamos (Greece)

Presented here are the deinotheriid dental and postcranial remains from the Late Miocene localities Pikermi and Halmyropotamos (Greece). The study and comparison of the available juvenile dental material from Pikermi with other relevant specimens from Europe showed that it belongs to the huge-sized Deinotherium proavum (= D. gigantissimum). Additionally, several postcranial specimens from Pikermi, as well as from Halmyropotamos, present deinotheriid features, which distinguish them from elephantoids, and permit their attribution also to D. proavum. This species is known from the Turolian and represents the terminal evolutionary stage of the European deinotheres. Its presence in Pikermi and Halmyropotamos is compatible with the middle Turolian (MN 12) age of the localities. Additionally, the taxonomy of European deinotheres is discussed, as well as the biostratigraphical and geographical distribution of D. proavum.

Equidae

Hipparionines represent the greatest part of the Nikiti 2 (NIK) fossil mammal collection. The determination of the material suggests the presence of four distinct species, two small, a medium and a large sized form. The small-sized forms attributed to Hipparion macedonicum, well known in northern Greece, and to H. sithonis nov. species. The weak canine fossa and the retraction of the narial opening above P2-P3 separate the latter species from H. macedonicum. Furthermore H. sithonis is slightly larger with more robust metapodials. The medium-sized NIK hipparion differs from other known medium-sized forms, representing the new species Hipparion philippus. Finally the large-sized form, represented in the studied material only by few specimens, has been assigned to H. cf. proboscideum. The morphological characters of the studied material and their stratigraphic range suggest an early Turolian to lower middle Turolian (MN 11–lower MN 12) age for the NIK fauna. Last, based on the hipparions found in NIK and the Eastern Mediterranean province in general, some hypotheses about the palaeobiogeography of the Aegean area and the phylogenetic relations among the hipparion species are discussed.

Hipparion macedonicum revisited: New data on evolution of hipparionine horses from the Late Miocene of Greece

The new expeditions to the Axios Valley (Macedonia, Greece) provided a new set of fossils from the various localities. Among the material collected from the Late Miocene hominoid bearing mammal locality Ravin de la Pluie (RPl) were some remains of Hipparion macedonicum, which was originally described from this locality. The most important is the skull and associated mandible, the first from the type locality. The new material is compared with the previously collected material of the taxon from the Vallesian and Turolian levels of Greece, as well as with corresponding material from Eurasia. The RPl skull is compared with the type skull of H. matthwei, a taxon, which several times has been referred as synonym to H. macedonicum. The comparison indicated several differences which distinguish the two species. Hipparion macedonicum has a continuous stratigraphic range from the Vallesian to middle Turolian and it is possibly present in the late Turolian. The comparison of the chronologically different samples of...

Fossil record and dynamics of Late Miocene small mammal faunas of the Vienna Basin and adjacent basins, Austria

The Late Miocene small mammal assemblages of the hinterland of Lake Pannon in Austrian Basins are represented by 99 species-level taxa and 30,400 specimens. The fossil-bearing localities can be grouped into eight intervals spanning about three million years from the Early Vallesian to the Middle Turolian. Each time slice is characterised by the occurrence and/or dominance of certain species. The retreat of Lake Pannon is reflected by a distinct diversification. This pattern may be a regional signal due to increasing habitat availability but may also be taphonomically biased due to a rather poor Earliest Vallesian record. Nevertheless, the overall community structure is quite stable throughout the Vallesian and no indication of a Vallesian Crisis can be detected. Instead, a moderate turnover occurs with the onset of the Turolian, reflected by the increasing abundance of xerophilic taxa.

New material of Chalicotheriidae (Perissodactyla, Mammalia) from the Late Miocene of Axios Valley, Macedonia (Greece) with the description of a new species

Chalicotheriids are rare in the late Miocene mammal localities of Axios Valley, Macedonia (Greece). The new campaign of excavations, since 1972, has provided some specimens, which are studied in this article. They are coming from two different localities. The late early Vallesian locality of Pentalophos 1 (PNT) has provided a skull and a mandible of an Ancylotherium. The morphological characters of the PNT material as the small size, the long snout, the shallow mandibular corpus, the strong cingulum in the teeth, the short tooth rows and the short M3/m3 indicate that it differs from the known Turolian species A. pentelicum and allow the erection of a new species, named Ancylotherium hellenicum n. sp., which can be used as a biostratigraphic marker of the Vallesian. The middle Turolian locality Prochoma 1 (PXM) has provided only one M3, which is determined to the chalicotheriine Anisodon macedonicus. This species was earlier described from the middle Turolian locality Vathylakkos 3 (VAT) and the late Turolian one of Dytiko 3 (DKO) of Axios Valley. The biogeography and biostratigraphy of the late Miocene chalicotheres of the Greco-Iranian Palaeoprovince (GRIP), as well as their palaeoecology are also discussed. The common chalicothere of GRIP is A. pentelicum, expanded from the Balkans to Afganistan and ranging stratigraphically from the early to the late Turolian. Chalicotherium goldfussi is certainly present in GRIP and it also ranges from the early to the late Turolian; its possible Vallesian occurrence needs confirmation. The other two late Miocene chalicotheres of GRIP A. macedonicus and Kalimantsia bulgarica are restricted to the Turolian of the Balkan Peninsula.

First occurrence of carnivore footprint with hyaenid affinities from the Late Miocene of Crete (Greece)

Nine terrestrial mammal localities of Miocene age based on body fossils have been reported from the island of Crete to date. A new locality where footprints of terrestrial mammals were exposed has recently been discovered in western Crete. Platýlakkos, the locality, is situated near the village Vouves, west of the city of Chania. The ichnofossils come from lacustrine deposits that belong to the Chatzi Formation. The most impressive finding is a clearly defined footprint that might be referred to a large-sized hyaenid. The footprint impression, a convex hyporelief on a sandstone slab, is a natural cast of a left manus where the traces of the interdigital pad and all four digital pads, as well as their respective claw marks- are clearly visible. Based on nearby marine deposits of the Chatzi Formation, the age of the fossiliferous layer with the footprints can be considered to be early to middle Turolian (MN11-MN12) in terms of mammalian biochronology. To date, there is no certain record of Miocene large-sized hyaenid footprints worldwide, and thus this isolated footprint might belong to a new ichno- taxon. However, in the absence of a trackway, the erection of a new ichnotaxon is avoided. The Platýlakkos findings are the first recorded case of fossil land mammal footprints from the Neogene of Crete, and Greece in general. They further the evidence for the presence of well-established terrestrial environments and faunas in the area of Crete during the Late Miocene.

A hominid tooth from Bulgaria: The last pre-human hominid of continental Europe

A hominid upper premolar was discovered in the Azmaka quarry, near Chirpan (Bulgaria). The associated fauna, especially the co-occurrence of Choerolophodon and Anancus among the proboscideans, and Cremohipparion matthewi and Hippotherium brachypus among the hipparions, constrains the age of the locality to the second half of the middle Turolian (ca. 7 Ma), making it the latest pre-human hominid of continental Europe and Asia Minor. The available morphological and metric data are more similar to those of Ouranopithecus from the Vallesian of Greece than to those of the early to middle Turolian hominids of Turkey and Georgia, but the time gap speaks against a direct phyletic link, and Turolian migration from the east cannot be rejected.

Şerefköy-2, a new Late Miocene mammal locality from the Yatağan Formation, Muğla, SW Turkey

Here we report on a new fossil locality, Şerefköy-2, from the Yatağan Basin of southwestern Turkey that preserves a well-sampled, abundant, and diverse mammal fauna. Indeed, after three field seasons, more than 1200 catalogued specimens representing 26 mammal species belonging to 14 genera make the Şerefköy-2 mammalian assemblage one of the richest Late Miocene fauna from Anatolia. Five hipparionines, six bovids, including the rare and enigmatic Urmiatherium rugosifrons and the presence of Pliohyrax graecus, strongly support affinities with Late Miocene faunas from Samos Island, Greece. Through a consideration of the identified material and the subsequent comparison with material from well-known Balkan and Anatolian faunas, a Middle Turolian (MN12) age for Şerefköy-2 is indicated.

A synopsis of the late Miocene Mammal Fauna of Samos Island, Aegean Sea, Greece

As one of the oldest known Eurasian fossil vertebrate localities, Samos late Miocene fauna attracted the interest of specialists by its richness and overall importance. Nevertheless, crucial taxonomical questions and chronological problems obscured its value. The detailed study of the local stratigraphy, the collection of new fossil material and its study, the revision of the old collections and the updated magneto-chronology of the fossiliferous deposits permited to re-discuss most of the problems in a special volume edited in 2009 by Koufos and Nagel and to provide a clearer and more precise idea about the Samos fauna and its age. A synopsis of this work is given here. The systematic study of the new collection (∼1200 identified specimens) allows the determination of 42 species from three fossil horizons, ranging from the upper part of early Turolian (MN11) to the end of middle Turolian (MN12). Taxonomic novelties are the presence of the carnivore genus Protictitherium found for the first time in Samos, the establishment of the new name Skoufotragus for Pachytragus Schlosser with the new species Skoufotragus zemalisorum, and the amended morphology of Pseudomeriones and Urmiatherium. Additionally six Hipparion and four Gazella species were recognized and a better morphometric distinction between Samotherium boissieri and Samotherium major was performed. This study also improved the correlation of the old fossiliferous sites with the new ones and with the local stratigraphy of the Mytilinii Basin, while precise ages have been obtained for the mammal localities. The new data together with the old collections indicate the presence in Samos of four chronologically successive mammal assemblages reflecting a “four stages-of-evolution” scheme. The Turolian palaeoenvironment of Samos is determined as an open bushland with thick grassy-floor of C3 graminoids with possible increase of the open and dry character from the beginning to the end of Middle Turolian. The Samos mammal faunas are palaeobiogeographically closer to the Asian ones than to those from the Greek mainland.