Biostratigraphy and biochronology of the Late Miocene of central Argentina: Evidence from rodents and taphonomy

Biostratigraphy and biochronology of the Late Miocene of central Argentina is analyzed through evolutionary patterns of octodontoid rodents. The studied faunas were found in continental sediments assigned to the Cerro Azul and Saldungaray formations (La Pampa and Buenos Aires provinces, respectively). Taphonomic attributes of recovered mammals, linked mostly to the development of paleosols or to predators’ activities, constraint the deposition lapse to short intervals. A biostratigraphic scheme based mainly on the anagenetic pattern of change in the octodontoid rodent lineage Chasichimys-Xenodontomys is proposed. Since there is no stratigraphic superposition among the studied levels, relationships among deposits are established through the polarity evidenced by the stage of evolution of chronomorphs of this lineage. Five new biozones for the Late Miocene of central Argentina are recognized. Evidence of other anagenetic sequences, and general trends of increasing hypsodonty in octodontoids, support this proposal. The results enhance the value of the “stage of evolution” concept as a correlation tool in biostratigraphic and biochronologic studies, mainly when applied through phyletic sequences.     

Early Miocene cricetids (Rodentia) from the Junggar basin (Xinjiang, China) and their biochronological implications

Among the 14 small mammal species from the early Miocene locality of the northern area of the Junggar basin (northern Xinjiang, China), four species are cricetids (Rodentia): unnamed species of Cricetodon and Eumyarion, and two new species, Karydomys debruijni nov. sp. and Megacricetodon beijiangensis nov. sp. Some aspects of the morphology of Cricetodon sp. are shared by Eucricetodon from the late Oligocene, suggesting that these specimens could be of intermediate form between Eucricetodon and Cricetodon. One tooth of Eumyarion sp. was found, making its determination uncertain, but its morphology is clearly differentiated from the one of Cricetodon sp. The species K. debruijni nov. sp. is established based on its primitive features compared to the species known in Europe and Anatolia, and its specific association of characters compared to Karydomys dzerzhinskii. M. beijiangensis nov. sp. shows many plesiomorphic features compared to the species already described in the middle Miocene of China. Based on both the whole assemblage of rodents and the species of cricetids, the biochronologic position and the age of the locality are discussed. The locality appears to be biochronologically very close to the fauna from the Chul’adyr Formation in Aktau Mountains, but we propose an age slightly older than the one proposed for this fauna, probably equivalent to the MN3 biozone in Europe.     

New Late Miocene murids from China (Mammalia, Rodentia)

New murids of Late Miocene (medial Baodean Chinese Mammal Unit) age from Inner Mongolia, northern China, and from Yunnan Province, southern China, are described. Hansdebruijnia perpusilla nov. sp. represents the earliest known and morphologically most primitive species of the genus, which is known from the latest Miocene of Europe and western Asia. The new species suggests an eastern Palaearctic origin of Hansdebruijnia. “Progonomys” yunnanensisQIU and STORCH, 1990 from Lufeng, Yunnan Province, is referred to Linomys nov. gen. New samples from Leilao, Yuanmou County, Yunnan Province, are included in this species, although this population is somewhat more primitive than that from Lufeng. Leilaomys zhudingi nov. gen. and sp. from Leilao shows a unique combination of apomorphic and plesiomorphic characters, which suggest an early divergence from the murid stem. The new findings indicate that our knowledge of the early radiation of murids in southeast Asia is still in its infancy.     

New materials of Dinocrocuta (Percrocutidae, Carnivora) from Lantian, Shaanxi Province, China, and remarks on Chinese Late Miocene biochronology

New materials from the middle part of the Bahe formation are described as Dinocrocuta gigantea. Review of the species reveals that it is derived in the evolutionary lineage of Dinocrocuta, and biochronologically later than Vallesian records from Turkey. The only possibly related Vallesian species from China is Crocuta gigantea xizangensis from Biru, Tibet, which may prove to be conspecific with D. senyureki. Based on the mammalian faunal sequence from Lantian, and with reference to Red Clay paleomagnetic data, the duration of D. gigantea in China should be later late Miocene, rather than the previously postulated early late Miocene (Vallesian equivalent) age.     

Vertebrate taphonomy in loess-palaeosol deposits: A case study from the late Miocene of central Argentina

This paper deals with the taphonomic analysis of the late Miocene bone assemblage from the Cerro Azul Formation at Telén (La Pampa Province, Argentina). The faunal assemblage was assigned to the Huayquerian mammal age (late Miocene). The fossiliferous section shows a homogeneous lithology, and is interpreted as a loess deposit with two similar and slightly developed palaeosols, classified as calcic vertisols. The studied sample comprises 5598 remains anatomically and taxonomically determined. They were collected from an area of about 48,000 m², appearing randomly distributed through the section and with low density. Most remains are small- to very small-sized, disarticulated, and very fragmented. Different taphonomical histories are inferred for microvertebrates and macromammals. The microvertebrate assemblage is interpreted as the result of predator activities. After a brief period of pre-burial exposure, remains were dispersed from the original depositional area. On the other hand, a natural and gradual death process is envisaged for macromammals, followed by a long period of exposure to weathering and dispersal by physical agents. Remains of both groups, once buried, suffered the diagenetic processes of the host rock. Consequently, the fossil assemblage from Telén would represent a condensed assemblage corresponding to two distinct time spans, i.e., the accumulation of microvertebrates took place in a short time interval whereas that of macromammals occurred over a longer period, coincident with the development of both soils.     

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.     

Prostrepsiceros and Protragelaphus (Artiodactyla,Mammalia) from the Late Miocene locality of Chomateri (Attica,Greece)

The Late Miocene fossiliferous locality of Chomateri is located close to the classic locality of Pikermi, but unfortunately its faunal context is not sufficiently known. Some fossil remains of spiral-horned antelopes from Chomateri are assigned to Prostrepsiceros rotundicornis and Protragelaphus skouzesi. The co-existence of P. rotundicornis and P. skouzesi is relatively rare in the Late Miocene, but well-documented in Pikermi. Comparisons of the Chomateri material support a middle Turolian age for the Chomateri locality and suggest a close chronological relation to the Pikermi fauna.     

Hypsosteiromys (Rodentia, Hystricognathi) from the Early Miocene of Patagonia (Argentina), the only Erethizontidae with a tendency to hypsodonty

Hypsosteiromys is the only New World porcupine that shows a tendency to hypsodonty. It is recorded exclusively from the Colhuehuapian Age (Early Miocene) of central Patagonia (Argentina). In addition to the type species, a second one, Hypsosteiromys nectus(AMEGHINO, 1902), is recognized, from the southern cliff of Colhuehuapi Lake (Chubut province). It differs from the type species in the lesser development of the anterolabial and posteroflexid notches of the m1-3, shorter p4 and dp4, and more slender incisors. Dental morphology suggests that the species of Hypsosteiromys lived in more open areas than most fossil and living Erethizontidae.     

Tropical freshwater teleosts from Miocene beds of eastern Patagonia, southern Argentina

Loricariid and pimelodid-like siluriforms—along with undetermined percomorph—are described for the first time from Patagonia. Vertebrate fossils, including fish, mammals and birds, were found in beds attributed to the top of the Puerto Madryn Formation. These levels supposedly corresponded to the “Rionegrense marino” of former authors. Mammals occurring in the site suggest a Huayquerian age for the fossiliferous beds. The base of the Huayquerian was dated at about 9 Ma and the top is younger than 6.5 Ma. Radioisotopic dating in the marine shell beds of the Puerto Madryn Formation ranges from 11 to 9 Ma. Consequently, the section described here appears to be younger than the typical Puerto Madryn Formation from which it is separated by an unconformity. The section is correlated with the type Río Negro Formation from northern Patagonia, which also includes Huayquerian fossils. Freshwater fishes were previously unknown in beds younger than the middle Miocene in southern South America. This is also the southernmost record of loricariid fishes. The association of aquatic continental and terrestrial vertebrates indicates for the first time in Península Valdés beds of freshwater origin. The evidence apported by fossils is also in agreement with global climate trends. The local extinction of loricariids in Patagonia possibly occurred much later than the time of deposition.     

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.     

Carnivores from the early/middle Miocene locality of Antonios (Chalkidiki, Macedonia, Greece)

The mammal locality of Antonios is one of the few known early-middle Miocene ones of Greece with large mammals. It is situated in Chalkidiki Peninsula (Macedonia, Greece) and includes both small and large mammals. The study of the carnivores found in Antonios indicates the presence of the following taxa: Proputorius cf. P. sansaniensis, Protictitherium gaillardi, Protictitherium cf. P. crassum, Percrocuta sp., Percrocutidae indet. and Pseusaelurus romieviensis. The material of each taxon is described, compared with other material from various Eurasian localities and determined. The species P. cf. sansaniensis and P. romieviensis are traced for first time in Eastern Mediterranean, while Percrocuta for first time in Greece. The presence of the other carnivoran taxa in the Antonios fauna is also interesting as they are recognized in older stratigraphic levels enriching our knowledge about their stratigraphic distribution. The carnivores of Antonios cannot offer significant data for the dating of the fauna but its age is discussed using the data coming from the carnivores as well as those from older studies of the micromammals and suoids.     

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”.     

Systematic revision of Siwalik Rhizomyidae (Rodentia)

Larges samples of fossil rhizomyids from the Siwalikmolasse deposits of the Potwar Plateau, Pakistan, enable reevaluation of fossil taxa and their evolutionary relationships. Siwalik Rhizomyidae constitute a basis for analysis of poorly represented rhizomyids from other areas in Asia and contribute to biochronological correlation. The extant Tachyoryctinae and Rhizomyinae diverged during the early history of the family and diagnoses of the subfamilies are amended to include extinct genera. Siwalik tachyoryctines include Kanisamys, Protachyoryctes, Rhizomyides, and Eicooryctes nov. gen.; rhizomyines are Brachyrhizomys and Anepsirhizomys nov. gen. Kanisamys, the earliest Siwalik rhizomyid, appeared by about 13 Ma and evolved slowly, displaying stasis in K. sivalensis. Brachyrhizomys evolved from Kanisamys by 9.5 Ma and radiated rapidly. B. nagrii displays rapid increase in size through time. About 7 Ma, Kanisamys and Brachyrhizomys became extinct locally and Protachyoryctes, Eicooryctes, and Rhizomyides appeared, perhaps in response to increasing aridity. After 5.5 Ma, rhizomyids became uncommon in the Potwar and Anepsirhizomys (3.0 Ma) constitutes the latest record of Rhizomyidae in Pakistan.     

A new elasmothere (Perissodactyla, Rhinocerotidae) from the late Miocene of the Linxia Basin in Gansu, China

Ningxiatherium euryrhinus sp. nov. is a relatively large elasmothere rhino from the Linxia Basin in northwestern China found in the early Bahean, which corresponds to the early Vallesian (MN 9), ca. 11.1 Ma. It is much larger than the extant Ceratotherium simum and has a single horn. Ningxiatherium is similar to the late Miocene Parelasmotherium from Gansu and Shanxi, but differs in having partially ossified nasal septum, terminal nasal horn boss, shallower nasal notch above the P3/P4 boundary, and much shorter premolars. N. euryrhinus sp. nov. differs from N. longirhinus from the late Miocene of Zhongning, Ningxia in northwestern China by its larger size, relatively wider nasals, shallower nasal notch, and sub-quadrangular occlusal surface of M3. In the Linxia Basin, the other early late Miocene (Bahean or Vallesian) elasmotheres are known from complete skulls or isolated teeth. They include two genera and three species: Parelasmotherium linxiaense, P. simplum, and Iranotherium morgani. They lack a nasal septum, but Ningxiatherium has one. The age of N. euryrhinus is early late Miocene based on direct association with biochronologic indicator taxa, such as Dinocrocuta gigantea, Hipparion dongxiangense, and Chilotherium primigenius. N. euryrhinus is more primitive than N. longirhinus sp. nov. in having a sub-quadrangular M3, a shallower nasal notch, and the presence of DP1 in adults.     

New material of Ouranopithecus macedoniensis from late Miocene of Macedonia (Greece) and study of its dental attrition

During the last five years our continued excavations in the known late Miocene mammal localities of Macedonia (Greece) provided several new specimens of the hominoid primate Ouranopithecus macedoniensis. This new material includes maxillary and mandibular remains and it is described and compared to the old material of Ouranopithecus in the present article. The material of Ouranopithecus from the three known localities “Ravin de la Pluie” (RPl), “Xirochori 1” (XIR) and “Nikiti 1” (NKT) includes a complete series of tooth rows representing all wearing stages. Thus, the study of the dental wear of Ouranopithecus upper and lower teeth is studied and compared to that of the recent hominoids Gorilla and Pan, as well as to Australopithecus afarensis. The latter species is well known by a series of tooth rows of different wearing stages. The canine’s attrition of Ouranopithecus has a more derived pattern than that of the recent hominoids (Gorilla and Pan) and less derived than A. afarensis. The p3 of Ouranopithecus has similar attrition to that of A. afarensis, the attrition of the molars in Ouranopithecus, A. afarensis and Pan follows a similar pattern, while in Gorilla it is different.     

An almost complete skeleton of Metailurus parvulus (Carnivora, Felidae) from the late Miocene of Kerassia (Northern Euboea, Greece)

We describe a partial skeleton of Metailurus parvulus from the Turolian site of Kerassia 1 (Northern Euboea, Greece). The material, which consists of a mandible, the anterior and posterior limb-bone elements, some sternal bones and some vertebrae, is the most complete known of this species. The dental material is compared to specimens from Pikermi and Chomateri (Greece), and China. The limb-bones available offer us the possibility to discuss the status of some previously described specimens from Pikermi. The limb proportions indicate that M. parvulus had elongated posterior limbs relative to the anterior ones, which reflects developed jumping skills. M. parvulus had moderately developed cursorial abilities, intermediate between open and closed habitat felids, and probably frequented primarily relatively open woodlands.     

Ancylotherium pentelicum (Perissodactyla, Mammalia) from the Upper Miocene of central and western Turkey

We report here new fossils of A. pentelicum from two Turolian localities of central and western Turkey, Pinaryaka and Salihpasalar (Kemikalan). The material, which consists of well preserved distal limb-bones, is compared with the few previously described specimens, mainly from Samos and Pikermi, but the observed differences should probably be attributed to individual variation. There is no evidence of evolution of the species during the Middle Turolian of the eastern Mediterranean, where it filled a narrow ecological niche, before its extinction.     

New data on late Miocene mammals of Kohfidisch, Austria

Cervids and bovids of the Early Turolian mammalian fauna of Kohfidisch (Austria) are reviewed. A new species, Micromeryx mirus sp. nov. (Moschidae) is described.