Norbert SCHMIDT-KITTLER, Hans de BRUIJN & Constantin DOUKAS:
General Introduction

The geological setting, sedimentology and biostratinomy of the new vertebrate locality Maramena (near Serres in Macedonia. Greece) is described. Its fossil content comprises plant remains, abundant molluscs, some reptiles and a rich mammalian fauna. Concluding from the mammalian assemblage the biochronological position of the locality is between the reference faunas of MN 13 and MN 14 which corresponds to the Turolian-Ruscinian boundary interval. The paleoenvironment of the site seems to have been a lake surrounded by dense vegetation and with more dry landscapes in the neighbourhood.

Christa FRANK:
The Malacofauna (Gastropoda et Bivalvia)

The study of the molluscan material of the vertebrate locality Maramena showed at least 20 species. The small shells of Limacacea could belong to more than one species. As far as it was possible the species were described morphologically. Their points of relationship are discussed with reference to the literature. Because of the fact that the authors had no comparable material from Northeastern Greece at her disposal and that number and state of preservation of most of the specimens seemed insufficient for a doubtless diagnosis, descriptions of new species are kept in reserve previously. Surely new is a species of Parmacella and a big species of the Helicoidea group. The high importance of molluscan thanatocoenoses for an analysis of environmental conditions is used to interpret the former habitat.

Annette RICHTER:
Lacertilia (Squamata, Reptilia)

Amongst the squamates, the Maramena vertebrate fauna also contains some lizard remains, most of which are small tooth-bearing fragments. So far, there are one single agamid maxillary, many dentaries from a not identifiable species of Lacerta, some unusual scincomorphan jaws here believed to be rather scincoid, and one questionable dentary of Ophisaurus. Additionally, many osteoderms of Ophisaurus could be found. The lack of gekkonids and chamaeleons as well as the presence of the to date unknown scincoid teeth make the Maramena faunal association special.

Zbigniew SZYNDLAR:
Serpentes (Squamata, Reptilia)

The locality Maramena yielded a snake material of 94 fragmentary vertebrae, which because of their uncomplete preservation can be determined on the family level only. Following to this, the subfamilies Colubrinae and Natricinae as well as the families Elapidae – with the genus Naja – and Viperidae – with the genus Vipera – can be recognized. Two precaudal vertebrae of Vipera certainly belong to the "oriental viper" group, which species are distinguished by their larger size. The members of this species-group are living today in Asia and Africa as well as in the easternmost part of Greece and the Cyclades, while in the Miocene and Pliocene (MN 3 to MN 16) they were widespread over most of Europe.

Constantin S. DOUKAS, Lars W. van den HOEK OSTENDE, Constantine D. THEOCHAROPOULOS & Jelle W. F. REUMER:
Insectivora (Erinaceidae, Talpidae, Soricidae, Mammalia)

In this paper the Insectivora from the lignite quarry of Maramena are described. A total of 11 species, belonging to the families Erinaceidae, Talpidae, and Soricidae is found. Three new species are described: Schizogalerix macedonica DOUKAS, Amblycoptus jessiae DOUKAS, and Deinsdorfia kerkhoffi REUMER. The fauna is indicative of a moist and forested environment. The insectivore assemblage has a rather Ruscinian character.

Ottmar KULLMER & Constantin DOUKAS:
The deciduous dentition of Mesopithecus pentelicus WAGNER (Primates, Mammalia)

The Maramena deposits contain tooth remains of Mesopithecus pentelicus WAGNER. The recovered permanent and deciduous teeth belong to several individuals. The deciduous dentition is compared with some modern cercopithecine monkeys. Especially in the deciduous front teeth a lot of features are quite similar between Mesopithecus and Colobus. Mesopithecus can be assigned to the subfamily Colobinae (see STRASSER & DELSON, 1987), which is supported by the morphological comparison of the milk dentition of several cercopithecids.

Norbert SCHMIDT-KITTLER:
Carnivora (Mammalia)

The new vertebrate locality of Maramena yielded a small carnivore fauna of six species, documented by jaw fragments, isolated teeth and tooth fragments. Of the recorded taxa two are described as new species. These are the mustelids Martes lefkonensis n.sp and Promeles macedonicus n.sp. No Turolian species can be found in the carnivore material, whereas two are known from the Ruscinian. The presence of a Lutra-species indicates open water, Promeles suggests forest vegetation and the occurence of a hyaenid shows that open landscapes cannot have been very far.

Hans de BRUIJN:
Sciuridae, Petauristidae and Eomyidae (Rodentia, Mammalia)

The association from Maramena contains one ground squirrel: Spermophilinus turolensis, one chipmunk: Tamias atsali, three flying squirrels: Hylopetes macedoniensis, Pliopetaurista dehneli and Miopetaurista thaleri and two eomyids: Leptodontomys catalaunicus and Keramidomys karpathicus. With the exception of Hylopetes macedoniensis and Tamias atsali all these species are known from (Upper)Miocene and Lower Pliocene deposits in Central and Western Europe. None of these species has been recorded from Asia.

Leptodontomys catalaunicus and Keramidomys karpathicus have extremely long stratigraphical ranges, but Pliopetaurista dehneli and Miopetaurrsta thaleri are restricted to the Late Turolian (MN 13) and Early Ruscinian (MN 14).

The flying squirrels and the eomyids suggest that the Maramena fauna lived in a wet tropical forest with dense undergrowth and tall trees.

Gudrun DAXNER-HÖCK:
Some Glirids and Cricetids from Maramena and other late Miocene localities in Northern Greece

Some Glirids and Cricetids from Maramena, Monasteri and Lefkon (Strimon Basin in Northern Greece) are described. The small mammal assemblages range from the late Vallesian / MN 10 (Lefkon) to the Turolian/Ruscinian transition = MN 13/14 (Monasteri and Maramena). Myomimus dehmi (de BRUIJN 1966) and Kowalskia sp. are recorded from Lefkon, Myomimus maritsensis de BRUIJN, DAWSON & MEIN 1970 from Monasteri and Myomimus maritsensis de BRUIJN, DAWSON & MEIN 1970, Glis cf. minor KOWALSKI 1956, Muscardinus pliocaenicus KOWALSKI 1963, Kowalskia browni DAXNER-HÖCK 1992, Allocricetus cf. ehiki SCHAUB 1930 and Hypsocricetus strimonis DAXNER-HÖCK 1992 are described from Maramena.

Gerhard STORCH & Thomas DAHLMANN:
Murinae (Rodentia, Mammalia)

About 2000 murine molars from Maramena, Strimon basin, Greece, are assigned to five species. Occitanomys neutrum is referred to a new subgenus, Hansdebruijnia n. subgen., and the new species Micromys cingulatus n.sp. is described. Apodemus gorafensis and A. cf. dominans represent distinct lineages within the heterogenous genus Apodemus. Rhagapodemus primaevus is the most primitive species of its genus. The Murinae suggest a latest Turolian (upper part of MN 13) age of the fauna.

Hans de BRUIJN:
Lagomorpha (Mammalia)

The lagomorphs from Maramena represent a species of Ochotonidae – Prolagus cf. sorbinii and a species of Leporidae – Alilepus turolensis. The type localities of these species, respectively Monte Castellaro (Italy) and El Arquillo (Spain), are both of Late Turolian (MN 13) = Messinian Age. The occurrence of these species at Maramena thus supports the age assigned to this locality on the basis of the regional geology and the evolutionary stage of many other groups of mammals.

Paul SONDAAR & Véra EISENMANN:
The Hipparions (Equidae, Perissodactyla, Mammalia)

Maramena has yielded a few fossils of Hipparion belonging to one rather small, slender species, with very hypsodont teeth and a very peculiar third metatarsal (MT III): The distal articular breadth of the bone is larger than the supra-articular breadth. This feature is unique among Old World hipparions and is known only in the Nannippus of the Blancan of North America.

Hypsodonty, proportions of the MT III and scarcity of the hipparion fossils relatively to the other ungulates, are all characters one can expect in the zone 5 of SEN et al. (1978) and point to a post Turolian age. Surface findings including one very large astragalus and one middle-sized distal MC III of a second species suggest a Ruscinian age.

Meinolf HELLMUND:
Suidae (Artiodactyla, Mammalia)

The fossil Suidae of the vertebrate locality Maramena (Macedonia, Greece) from the Turolian-Ruscinian Boundary (MN 13/14) are described and compared with relevant European taxa of similar stratigraphic range like Eumaiochoerus HÜRZELER 1982, Microstonyx PILGRIM 1926 and Korynochoerus SCHMIDT-KITTLER 1971. The new specimens were attributed to the type species Korynochoerus palaeochoerus (KAUP 1833). K. palaeochoerus shows a characteristic sexual dimorphism concerning canine sup./inf. morphology; the alveolarcristae above C sup. being weak and only scarcely protruding laterally are assigned to female individuals, male individuals were not found in Maramena. Korynochoerus palaeochoerus obviously is a temporary extensive species, known from the Lower Vallesian (MN 9) to the Turolian-Ruscinian boundary (MN 13/14) of Europe.

Beatriz AZANZA:
Cervidae (Artiodactyla, Mammalia)

From the vertebrate locality Maramena a new three-tined deer, Pliocervus graecus n.sp., is described on the basis of antlers and teeth remains. The antlers from Pikermi and Halmyropotamus hitherto referred to Pliocervus pentelici (GAUDRY 1862) are also attributed to this new species. The dentition for P. graecus is rather evolved and corresponds in its dimensions to "Dremotherium" sp. from Pikermi. The species "Dremotherium" pentelici is retained to the small dentition from Pikermi.

Meike KÖHLER, Salvador MOYA-SOLA & Jorge MORALES:
Bovidae and Giraffidae (Artiodactyla, Mammalia)

Cranial and postcranial remains of seven bovids and one giraffid are described from the greek locality Maramena. The bovid fauna is composed of three Boselaphines, two Antilopines, one caprine and one indetermined bovid. The caprine form is described as a new genus and species, Norbertia hellenica n.g. n.sp. The giraffid is the large Samotherium. The fauna is mainly typical Upper Turolian, but the presence of two very modern mountain adapted forms (Norbertia and Bov. indet.) is surprising. Thus, for first time the appearence of a real member of the Caprinae is reported from that early mammal stage (MN 13/14).