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.     

New material of Carnivora (Mammalia) from the Late Miocene of Axios Valley,Macedonia, Greece

A new collection of carnivores from the Late Miocene deposits of Axios Valley, Macedonia, Greece was collected in 2004–2009 at the localities named Xirochori 1 (XIR) and Ravin de la Pluie (RPl), dated to the Late Vallesian (MN 10), as well as at Ravin des Zouaves 5 (RZO), dated to the Early Turolian (MN 11). The studied material is described and compared morphologically and biometrically with other materials from Greece and its neighboring area. The following species are determined: XIR: Dinocrocuta sp.; RPl: Eomellivora wimani, Adcrocuta eximia leptoryncha, ?Hyaenictis sp., Metailurus parvulus; RZO: Plioviverrops orbignyi, Machairodus giganteus. This collection is interesting because it includes the upper deciduous dentition of A. eximia and Dinocrocuta, which are only known from limited material until now, especially the latter taxon. E. wimani is recognized for the first time in Greece and the Balkans, while M. parvulus is identified for the first time in the Vallesian of Greece. The possible presence of Hyaenictis in RPl is the first indication of its presence in the Vallesian. The guild structure diagrams of the carnivore fauna from the three studied localities provide some indications of their palaeoenvironment, which fits quite well with the results of previous studies on the palaeoenvironments of these localities and of the wider Eastern Mediterranean region.     

Metailurus major Zdansky, 1924 (Carnivora,Mammalia) from the classical locality of Pikermi (Attica,Greece)

The species of Metailurus major has a large geographical extension and is known from localities spread out from West Europe to China. In Greece it is known from the Late Miocene locality of Halmyropotamos in Euboea, while some authors mention it also in Pikermi and Samos in faunal lists only. Two specimens from Pikermi prove its existence in this classical locality.     

<Emphasis Type="Italic">Mesopithecus</Emphasis> (Primates,Cercopithecidae) from the turolian locality of Vathylakkos 2 (Macedonia,Greece)

Two skulls, two mandibles and some postcranials ofMesopithecus from the locality “Vathylakkos 2” of Axios Valley (Macedonia, Greece) enlarge our knowledge about the genus and its differentiation. The new material together with an old skull from the same locality is compared with the Pikermi, “Ravin des Zouaves 5” (Axios Valley), and Maramena (Serres basin) samples. It is concluded that the VathylakkosMesopithecus resemblesM. pentelicus from Pikermi, as wellM. delsoni from “Ravin des Zouaves 5” and it is referred asMesopithecus sp. aff.M. pentelicus. The biochronological age of “Vathylakkos 2” has been considered as MN 12, while recent magnetostratigraphic data indicate an age of about 7.5 Ma. This age confirms the position of the VathylakkosMesopithecus between Pikermi and “Ravin des Zouaves 5”. Some dental indices compared with those of the RecentCercopithecus suggest that the Vathylakkos sample is monospecific.      

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.     

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.     

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.     

Merychippus (Mammalia, Perissodactyla, Equidae) from the Middle Miocene of state of Oaxaca, southeastern Mexico

Mexican material referable to Merychippus from two localities in eastern Oaxaca was described first nearly 50 years ago. Subsequent work there and in Central Oaxaca, spanning some 30 years, has allowed to establish the detail stratigraphy in both regions, and assembled a collection of merychippine material from the Matatlán (Central Oaxaca) and El Camarón (eastern Oaxaca) Formations, both K-Ar dated ~15 Ma (late early Barstovian). Detailed taxonomic analysis of this collection indicate the presence of two subhypsodont horse species referable to “Merychippus” cf. “M.” primus and “M.” cf. “M.” sejunctus in both regions. These records document the coexistence in tropical southern North America of basal and hipparionine affinity merychippine grade species, and provide a glimpse in to the diversity of subhypsodont equids in this region.     

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.     

Juvenile ornithopod (Dinosauria: Rhabdodontidae) remains from the Upper Cretaceous (Lower Campanian, Gosau Group) of Muthmannsdorf (Lower Austria)

The fragmentary remains of a juvenile rhabdodontid ornithopod from the Coal-bearing Complex of the Gosau Group (Lower Campanian, Grünbach syncline) at Muthmannsdorf near Wiener Neustadt, Lower Austria are revised. The material, probably belonging to a single individual, includes a right dentary (lectotype of Iguanodon suessi Bunzel, 1871, designated herein), teeth, a fragmentary parietal, fragments of scapula, ?radius, femur, tibia, two vertebrae (lost) and a manual ungual.The lectotype dentary does not provide clear autapomorphies or sufficient diagnostic features to determine its position within the Rhabdodontidae at generic level. By this “Iguanodon suessi” Bunzel, 1871 and the genus “Mochlodon” Seeley, 1881, to which it was latter referred as type species, cannot be characterized sufficiently by differential diagnosis and these are best considered nomina dubia. Based upon combined character comparisons (mainly postcranial features) the Muthmannsdorf ornithopod is referred herein to Zalmoxes Weishampel, Jianu, Csiki and Norman, 2003, a genus so far known from the late Maastrichtian of Romania. It probably but not evidently represents a yet unnamed species, most closely related to Zalmoxes shqiperorum Weishampel, Jianu, Csiki and Norman, 2003. At the present state of knowledge the Austrian material is not further diagnostic at the species level and kept in open nomenclature as Zalmoxes sp.     

First giraffid skulls (Bohlinia attica) from the late Miocene Maragheh fauna,Northwest Iran

An almost complete skull and a second partial skull of Bohlinia attica (Artiodactyla: Giraffidae) from the late Miocene of Maragheh in northwestern Iran is described along with a complete upper dentition from Samos in Greece. These specimens enrich what is known of this species. The skull has massive bent ossicones. The braincase is horizontal to the face in lateral view. The upper premolars have strongly curved styles. The metapodials of this species are elongate with a deep plantar trough. The braincase is short and the occipital does not extend caudally. The type material of Bohlinia is from Pikermi but this taxon is also known from localities in F.Y.R.O. Macedonia, Greece, Turkey, Bulgaria, Iraq, and Iran. The new specimens are similar to others except that the ossicone terminates in a knob and the palatine choanae are positioned caudally. This taxon along with Honanotherium, which is most similar but with shorter metapodials and simpler premolars, can be placed in the subfamily Bohlininae.     

Morphometric and molecular characterisation of specimens of Lepidapedon Stafford, 1904 (Digenea: Lepidapedidae) from the deep-sea fish Mora moro (Risso) (Teleostei: Moridae) in the western Mediterranean

In a study of the parasites of the deep-sea fish Mora moro (Risso) (Gadiformes: Moridae) off the Mediterranean coasts of Catalonia and the Balearic Islands (Spain), we were able to distinguish two morphs of specimens belonging to Lepidapedon Stafford, 1904 (Digenea: Lepidapedidae). This material is herein described and illustrated. Comparative sequence analyses using partial mitochondrial nad1 sequences revealed that the material assigned to one of these morphs can be considered conspecific with the material identified as Lepidapedon desclersae Bray & Gibson, 1995 from the same host. However, the published nad1 sequence for L. desclersae was generated from a specimen ex M. moro from the North East Atlantic. Examination of the voucher specimens associated with this sequence revealed that both the North East Atlantic and the Mediterranean specimens ex M. moro differ from L. desclersae as described from its type-host, Lepidion eques (Günther), in the anterior extent of the vitelline fields which is further posterior, reaching only to the posterior margin of the external seminal vesicle in L. desclersae, versus being at the mid-level of this organ and reaching the posterior margin of the ventral sucker. Therefore, we have tentatively assigned the material characterised here, both morphologically and molecularly as Lepidapedon sp. Acquisition of additional sequences for both nad1 mitochondrial and 28S rRNA genes of L. desclersae from material ex Lepidion spp. is required in order to determine whether the observed morphometric variation reflects host-related or inter-specific differences. The second morph of Lepidapedon from M. moro is described and distinguished on morphometric grounds, such as the position of the most anterior vitelline follicles, which reach to the anterior margin of the ventral sucker. Its identity is commented upon, but, in view of the fact that there were few specimens and no molecular data available, it is not named.     

Calcaneal features of the Greek Miocene primate Mesopithecus pentelicus (Cercopithecoidea: Colobinae)

This paper investigates substrate preferences of the Greek Colobine Mesopithecus pentelicus WAGNER, from the Miocene of Pikermi, by examining selected functional features of the calcaneus that distinguish between arboreal and terrestrial Cercopithecidae. Mesopithecus possesses a relatively long proximal calcaneal region associated with a slightly low and wide surface for the insertion of m. triceps surae. These features approximate that of semi-terrestrial Cercopithecidae and would suggest terrestrial cursorial activities. On the other hand, the relatively long and narrow proximal calcaneo-astragalar facet, similar to that of most arboreal species, would provide ampler subtalar movements. The mosaic of these features implies a semi-terrestrial way of life and conforms to the savanna-woodland paleoenvironment of Pikermi, Greece. These features appear to be well associated with foot function and change of habitus and are used to examine scenarios of the evolutionary history of the Colobinae.     

Marine reptiles (Thalattosuchia) from the Early Jurassic of Lombardy (northern Italy)

The fossil remains of two small reptiles recently discovered in the Sogno Formation (Lower Toarcian) near Cesana Brianza (Lecco Province), represent the first mesoeucrocodylians reported for Lombardy and some of the few Jurassic reptiles from Italy. Due to the absence of diagnostic skeletal elements (the skulls are lacking), it is not possible to refer the new specimens at genus level with confidence. Although the well developed dermal armour would characterise Toarcian thalattosuchians of the genera Steneosaurus (Teleosauridae) and Pelagosaurus (Metriorhynchidae), the peculiar morphology of the osteoderms allow to tentatively refer the remains to the latter taxon (cf. Pelagosaurus sp.). The small size, along with the opening of the neurocentral vertebral sutures and, possibly, the non sutured caudal pleurapophyses, indicate that the specimens were morphologically immature at death. These “marine crocodiles” confirm the affinities between the fauna of the Calcare di Sogno Formation and coeval outcrops of central Europe that also share the presence of similar fishes and crustaceans.     

Plio-Pleistocene evolution of the genus Castor (Rodentia, Mammalia) in Europe: C. fiber plicidens of Pietrafitta (Perugia, Central Italy)

In the lignite sediments of Pietrafitta (Tiberino Basin, Umbria, Central Italy), a rich fossil assemblage of vertebrate, invertebrate and plant remains belonging to the Farneta Faunal Unit (Late Villafranchian, Early Pleistocene) was collected. Among them fossil beaver cranial remains are relatively abundant. Western-Central Europe Villafranchian beaver remains were assigned to C. plicidens by some authors because molar occlusal surface patterns show complex enamel crenulations. Several recent authors have classified them as C. fiber while analysing other morphological patterns. Our samples have been compared to Plio-Pleistocene fossil remains and to living European populations of the genus Castor. New morphometric parameters on molar occlusal surfaces have been defined and statistical analyses (One-Way MANOVA, Principal Component Analysis and Canonical Discriminant Analysis) have been performed on them. The results point out a subspecific separation between the Late Villafranchian beaver of Pietrafitta, Quarata and San Giovanni in Valdarno (Umbria and Tuscany) and C. fiber populations. St. Vallier (France) Late Pliocene and Mosbach 2 (Germany) Middle Pleistocene beavers classified respectively by Viret and Friant as C. plicidens, show a C. f. fiber molar teeth pattern. Therefore, C. fiber plicidens did not occur in Central-Western Europe and this subspecific name may be used only for the local populations of Valdarno and Tiberino Basin (San Giovanni in Valdarno, Quarata, Pietrafitta and a few localities of the same area), at that time peripheral populations, probably semi-isolated during the Late Villafranchian, and at the southern limit of the geographic range of C. fiber.     

Additional specimens of Hyracoidea (Mammalia) from the Early and Middle Miocene of Kenya

We describe unpublished material of the poorly known hyracoid, Brachyhyrax aequatorialis from the Early Miocene of Songhor and Koru, Kenya, on the basis of specimens stored in the Community Museums of Kenya, the National Museums of Kenya and the Natural History Museum, London. As a result, we added 17 specimens to the hypodigm of this species which was previously known from only seven specimens. In addition, we describe further material of another hyracoid, Afrohyrax championi, from the Early Miocene of Mfwangano and the Middle Miocene of Kipsaraman. The chronological distributions of the two hyracoids do not overlap. Brachyhyrax appears to have lived in forest environments, a suggestion supported by its brachyodont dentition and the associated molluscan fauna, whereas Afrohyrax seems to have lived in more open wooded habitats which agree with its slightly more hypsodont dentition and the cursorial postcranial skeleton as well as with the land snails that occur with it.     

Evidence for the longest stratigraphic range of a post-Triassic Ichthyosaur: a Leptonectes tenuirostris from the Pliensbachian (Lower Jurassic) of Switzerland

A three-dimensional well-preserved ichthyosaur skull and parts of the postcranial skeleton are attributed to the species Leptonectes tenuirostris (Conybeare, 1822). It was found vertically embedded in Pliensbachian deposits representing three successive biozones (ibex to margaritatus Zone). The find is dated as early Late Pliensbachian (margaritatus Zone) by a rich ammonite and ostracod fauna. It is the first record of the genus Leptonectes from Switzerland and from the Late Pliensbachian. It is so far the best preserved and most complete ichthyosaur from this time interval worldwide. With diagnostic specimens known from the Rhaetian (Late Keuper) up to the early Late Pliensbachian (Middle Liassic), L. tenuirostris (Conybeare, 1822) has the most extensive stratigraphic range documented for any post-Triassic ichthyosaur so far.     

Investigation of the ultrastructure of Dendrocoelum constrictum (Platyhelminthes,Tricladida) spermatogenesis and mature spermatozoa

To add to our understanding of dendrocoelid spermatozoa and to describe additional phylogenetic characters, the ultrastructure of the testis was investigated in the subterranean freshwater planarian Dendrocoelum constrictum. This is the first study investigating spermatogenesis and spermatozoon ultrastructure in a subterranean freshwater planarian species. We found that the basic structure of spermatozoa in D. constrictum is similar to that of other Tricladida that have been studied previously. In fact, D. constrictum spermatozoa possess an elongated nucleus, one giant mitochondrion, and two subterminal flagella with a 9 + ‘1’ pattern. The flagella emerge together from one side of the spermatozoon. However, D. constrictum has some characteristics that have not yet been described for other freshwater planarians. In fact, the number of cortical microtubules reaches the maximum number in the anterior and middle part of region I, and then decrease until they disappear towards the posterior extremity of the spermatozoon. The extreme tip of the anterior region of the spermatozoon exhibits a specific external ornamentation of the plasma membrane.