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.     

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.     

A ctenodactylid rodent (Mammalia: Rodentia) from the Middle Miocene of Chios Island (Greece)

Field work carried out in 1991 and 1993 at the Middle Miocene locality of Thymiana (Chios Island, Greece), produced much rodent material including a number of ctenodactylid dental specimens. They represent a single taxon whose upper and lower cheek tooth morphology clearly differs from Prosayimys flynni and all Sayimys species known so far, except for Sayimys intermedius. The ctenodactylid teeth from the Middle Miocene of Chios are identified as pertaining to the latter species, despite minor differences from the Pakistani and Saudi Arabian representatives of S. intermedius.     

A new lorisid humerus from the early miocene of Uganda

An early Miocene (18–20 mya) distal humerus from Napak, Uganda, is the only element of the forelimb to be allocated to African Lorisinae. Its loris-like features distinguish it from other East African material that more closely resembles galagids. Tracing the evolution of the lorisoid body has been a difficult task due to the lack of good fossil material. Here, we add a single specimen to this poor record. This humerus helps to document the beginning of extant lorisid locomotor specialization and function.     

Analysis of samples of Hipparion(Mammalia,Perissodactyla) from the Concud localities,Turolian of Spain

Samples, mainly of isolated teeth, of Hipparion concudensePirlot from the Turolian fossil sites “Concud”. Cerro de la Garita, Barranco de las Calaveras, Concud III, Masia del Barbo, and Los Mansuetos, all in the Calatayud-Teruel area, Spain, are analysed and compared. The teeth from “Concud”, Cerro, and Masia are similar and larger than those from Barranco, Concud III, and Los Mansuetos, which between them are similar. A possible trend towards decrease in size is indicated. The stratigraphic position of these sites is briefly discussed.     

Locomotor evolution of <Emphasis Type=Italic>Mesopithecus</Emphasis> (Primates: Colobinae) from Greece: evidence from selected astragalar characters

This paper reports our investigations into functional aspects of the astragalus of four samples of the genus Mesopithecus from Greece. More particularly, it aims to infer substrate preferences of M. delsoni/pentelicus from the Middle Turolian site of Perivolaki (central Greece), M. pentelicus from the late Middle Turolian site of Pikermi (southern Greece), and M. cf. pentelicus and M. cf. monspessulanus from the Late Turolian site of Dytiko (northern Greece). For these purposes, selected astragalar functional features, such as trochlea wedging, proximal facet curvature, and head rotation were expressed as linear measurements on both fossil and selected extant colobines. The size-adjusted measurements were used for univariate comparisons as well as a multivariate principal components analysis. Both approaches revealed that the selected characters were able to discriminate between extant arboreal and semiterrestrial colobines, but all fossil forms presented mosaic morphology. Thus, the oldest representative, M. delsoni/pentelicus was reconstructed as mainly semiterrestrial. On the other hand, the astragalus of M. pentelicus appeared to reflect semiterrestrial habits with a moderate adaptation to arboreality. Similar habitus reconstruction was allocated to the more recent M. cf. pentelicus, whereas the sympatric and synchronous M. cf. monspessulanus showed semiterrestrial adaptations with a slight preference of terrestrial substrates. The results mainly conform to paleoenvironmental reconstructions of the fossiliferous localities and denote that Mesopithecus was mainly a semiterrestrial radiation throughout its evolutionary history, with differential rates of use between arboreal and terrestrial substrates. These adaptations could have promoted the dispersal of the genus throughout Eurasia during the latest Miocene and Early Pliocene.     

The deciduous lower dentition of Ouranopithecus macedoniensis (Primates, Hominoidea) from the late Miocene deposits of Macedonia, Greece

Two mandibular fragments with associated milk teeth assigned to the late Miocene hominoid primate Ouranopithecus macedoniensis are analyzed. The fossils, which belong to a single individual, were found in the Vallesian locality of "Ravin de la Pluie" of the Axios Valley (Macedonia, Greece). The material is described here and compared with extant and extinct hominoids, allowing assessment of the evolutionary trends in the deciduous lower dentition within the Hominoidea. Hylobatids represent the more primitive pattern. Gorilla is slightly more derived than hylobatids, but less derived than Pongo and Pan, the latter being the most derived. With relatively smaller deciduous canines and more molarized deciduous premolars, Ouranopithecus is more derived than both Pan and Gorilla. Among the fossil hominoids, Proconsul, representing the primitive condition, has a very simple dp(3)and a dp(4)that has a trigonid that is taller than the talonid and which lacks a hypoconulid. Griphopithecus is more derived than Proconsul in having a dp(4) with a lower trigonid, a hypoconulid, and a less oblique cristid obliqua. Australopithecus and Paranthropus possess a similar morphology to that of Homo, while Ardipithecus appears to be more primitive than the latter genera. Ouranopithecus has a more derived lower milk dentition than Proconsul and Griphopithecus, but less derived than Australopithecus and Paranthropus. The comparison of the lower milk dentition of Ouranopithecus confirms our previous conclusions suggesting that this fossil hominoid shares derived characters with Australopithecus and Homo.     

Primate numts and reticulate evolution of capped and golden leaf monkeys (Primates: Colobinae)

A recent phylogenetic study of langurs and leaf monkeys of South Asia suggested a reticulate evolution of capped and golden leaf monkeys through ancient hybridization between Semnopithecus and Trachypithecus. To test this hybridization scenario, I analysed nuclear copies of the mitochondrial cytochrome b gene (numts) from capped, golden and Phayre’s leaf monkeys. These numts were aligned with mitochondrial cytochrome b sequences of various species belonging to the genera Semnopithecus and Trachypithecus. In the phylogenetic tree derived from this alignment, the numts fell into three distinct clades (A, B and C) suggesting three independent integration events. Clade A was basal to Semnopithecus, and clades B and C were basal to Trachypithecus. Among the numts in clades A and C were sequences derived from species not represented in their respective sister mitochondrial groups. This unusual placement of certain numts is taken as additional support for the hybridization scenario. Based on the molecular dating of these integration events, hybridization is estimated to have occurred around 7.1 to 3.4 million years ago. Capped and golden leaf monkeys might have to be assigned to a new genus to reconcile their unique evolutionary history. Additionally, northeast India appears to be a ‘hot spot’ for lineages that might have evolved through reticulate evolution.     

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.     

Molecular phylogeny and biogeography of langurs and leaf monkeys of South Asia (Primates: Colobinae)

The two recently proposed taxonomies of the langurs and leaf monkeys (Subfamily Colobinae) provide different implications to our understanding of the evolution of Nilgiri and purple-faced langurs. Groves (2001) [Groves, C.P., 2001. Primate Taxonomy. Smithsonian Institute Press, Washington], placed Nilgiri and purple-faced langurs in the genus Trachypithecus, thereby suggesting disjunct distribution of the genus Trachypithecus. [Brandon-Jones, D., Eudey, A.A., Geissmann, T., Groves, C.P., Melnick, D.J., Morales, J.C., Shekelle, M., Stewart, C.-B., 2003. Asian primate classification. Int. J. Primatol. 25, 97-162] placed these langurs in the genus Semnopithecus, which suggests convergence of morphological characters in Nilgiri and purple-faced langurs with Trachypithecus. To test these scenarios, we sequenced and analyzed the mitochondrial cytochrome b gene and two nuclear DNA-encoded genes, lysozyme and protamine P1, from a variety of colobine species. All three markers support the clustering of Nilgiri and purple-faced langurs with Hanuman langur (Semnopithecus), while leaf monkeys of Southeast Asian (Trachypithecus) form a distinct clade. The phylogenetic position of capped and golden leaf monkeys is still unresolved. It is likely that this species group might have evolved due to past hybridization between Semnopithecus and Trachypithecus clades.     

Remains of Hipparion (Equidae,Perissodactyla) from Puente Minero (Teruel Province,Spain) and their implications for the systematics of the Turolian Hipparionini

Abstract: Morphological and biometrical variability of the equid Hipparion from Puente Minero and other localities from the Teruel Basin, Spain (MN10–MN13), is analysed. Three species of Hipparion are recognized in Puente Minero, H. laromae, H. matthewi and Hipparion sp. cf. H. longipes, through comparison with other Spanish and Eurasian species (Samos, Greece; Höwenegg, Germany; Pavlodar, Kazakhstan, small and large forms; Akka?da?i, Turkey; and the following Spanish sites, La Roma 2, Los Aljezares, Concud, Milagros, Las Casiones and Venta de Moro). The majority of the Hipparion remains from Puente Minero and determined to be Hipparion laromae based on their size and morphology. This interpretation was confirmed by bivariate and multivariate analyses. These results suggest a Eurasian immigration of different Hipparion species into the Teruel Basin.     

<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.      

A new stage in the evolution of the mole rats (Rodentia,Spalacinae) from the Early Miocene of northern Greece

The cheek teeth of Debruijnia kostakii n. sp. from the Early Miocene of Karydia are only slightly smaller than those of the largest extant species of the Spalacinae (Spalax giganteus), but show characteristics that are reminiscent of its Paleogene cricetid ancestor. The peculiar combination of large size and primitive dental morphology of this species bridges the gap between Debruijnia and Pliospalax and thereby supports the hypothesis that these genera form the core of the complex evolutionary history leading to extant Spalax.     

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”.     

Poor but classic: The squamate fauna from the late Miocene of Pikermi,near Athens,Greece

I here describe new fossil remains of lizards and snakes from the late Miocene of Pikermi, near Athens, Greece. This classical locality, known since at least the middle of the 19th century, has yielded over the decades abundant mammal fossils, whereas squamate remains have been confined to early discoveries of Varanus remains during the 19th century. The first specimen among these varanid fossils, a large vertebra described by Albert Gaudry during the 1860s, represents also one of the earliest discoveries of a fossil monitor lizard worldwide. Nevertheless, this historically important specimen has only been briefly described and its so far only available images were the original lithographs and subsequent sketches of two only of its views. I here describe this specimen in detail and provide photographs of all its standard views. The additional squamate remains from the Pikermi area are fragmentary. Nevertheless, they document the presence in Pikermi of other lizard groups, such as lacertids and probable anguines, while snakes are described for the first time. Among the snake fossils, probable erycines and colubrids are identified; if indeed an erycine, this would be the oldest occurrence of that group in the Eastern Mediterranean, thus extending back its stratigraphic range in the area by several million years.     

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.     

The radiation of red colobus monkeys (Primates,Colobinae): morphological evolution in a clade of endangered African primates

Red colobus monkeys are a group of African monkeys that include some of the most endangered primate populations. Despite urgently needing to understand the importance of particular populations for preserving the biodiversity of this lineage, their evolutionary relationships remain poorly understood, and their taxonomy is unstable, and often enigmatic. Data on behaviour, ecology, genetics, and morphology are thus strongly needed to address taxonomic issues that are not only relevant for primatologists, but also for conservation biologists. In this study, we investigated the morphological diversity and evolution of red colobus by examining the cranial variation of 369 individuals from most living populations. Crania were measured using a set of 64 anatomical landmarks, and were analysed using geometric morphometric methods for the study of three‐dimensional landmark coordinates. We found significant differences among most of the populations traditionally described on the basis of pelage colour and geographic distribution. However, differences tended to be smaller within biogeographic assemblages, which might be related to mountain refugia during periods of forest contraction in the Pleistocene.We also found a tendency towards large taxonomic distances, which suggested that populations might have originated earlier than has been traditionally thought, a result congruent with a recent molecular phylogenetic analysis. However, the distinctive forms of East African relict populations might be related to an acceleration of morphological evolution in small peripheral isolates, under strong selective pressures. This indicates that small and isolated populations, which are also the most endangered ones, might indeed be unique representatives of the red colobus radiation, and hence contribute to its biodiversity significantly. However, in‐ depth morphological studies of red colobus, particularly those in peripheral populations that tend to be rare in the wild, as well as in museum collections, is hampered by a paucity of data. In these cases, populations might be extinct before primatologists and conservationsts can even appreciate what was lost.     

Functional aspects of primate pelvic structure: A multivariate approach

This study aims to clarify the relationship of primate bony pelvic structure to locomotor habit. As with most of the postcranial skeleton, the pelvic bones of species within the Ceboidea and the Cercopithecoidea are remarkably similar visually except for variations in size. Yet there are substantial differences in locomotor pattern between the species in these taxa. I performed canonical analyses on a sample of 17 pelvic variables describing 22 primate species of the Ceboidea, the Cercopithecoidea, and the Hominoidea to discover which variables were significant in separating them into groups. In both analyses there was good separation of major taxa and additional separation of groups that differed in locomotor habit. The separation of colobine from cercopithecine monkeys was particularly consistent. In the analysis, including all 22 species, the variables given particular weight by the canonical analysis were the same as those traditionally used by anatomists for the same purpose. Specifically, breadth of the ischial tuberosity (reflecting presence or absence of ischial callosities) separated the Old from the New World monkeys. Breadth of the iliac tuberosity, in which man and to some extent other hominoids differ from other primates, and ilium height, in which man differs from other primates, were significant. Sagittal diameter of the pelvis was also substantially weighted. Having established that the technique would select variables of anatomical significance, the same method was applied to a study of monkeys only where the characteristics that differ between groups are not well established. Breadth of the ischial tuberosity was again important in separating the Ceboidea from the Cercopithecoidea. Discrimination of locomotor groups within these large divisions was brought about mainly by ischial length and the sagittal diameter of the pelvis. In studying these variables and their relationship to size in greater detail, it was found that among cercopithecoid monkeys, the colobines showed relatively lower values than did cercopithecines for both these dimensions. Atelines showed low values for ischial length but high values for the sagittal pelvic diameter. Biomechanical explanations of these observations are suggested.