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.     

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.     

<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.      

Mitochondrial and nuclear markers suggest Hanuman langur (Primates: Colobinae) polyphyly: Implications for their species status

Recent molecular studies on langurs of the Indian subcontinent suggest that the widely-distributed and morphologically variable Hanuman langurs (Semnopithecus entellus) are polyphyletic with respect to Nilgiri and purple-faced langurs. To further investigate this scenario, we have analyzed additional sequences of mitochondrial cytochrome b as well as nuclear protamine P1 genes from these species. The results confirm Hanuman langur polyphyly in the mitochondrial tree and the nuclear markers suggest that the Hanuman langurs share protamine P1 alleles with Nilgiri and purple-faced langurs. We recommend provisional splitting of the so-called Hanuman langurs into three species such that the taxonomy is consistent with their evolutionary relationships.     

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”.     

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.     

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.     

A multivariate analysis of Hyospathe (Palmae)

Previous systematic treatments of the neotropical palm genus Hyospathe have recognized from two to 18 species. An explicit, quantitative, repeatable sequence of operations for delimiting and testing groups of specimens and applying species concepts is carried out. Multivariate statistical analysis of morphological data is used to delimit and test groups of specimens. Cluster analysis is used to distinguish between characters and traits. Analysis of qualitative and quantitative characters reveals six groups of specimens, and the Phylogenetic Species Concept is applied to these groups. Two species, H. peruviana Henderson and H. frontinensis Henderson, are described as new. One of the specimen groups is large and widespread, and six geographically separate subgroups can be recognized within it. These subgroups can be distinguished by one or more significantly different quantitative characters. A Phylogenetic Subspecies Concept is applied to these subgroups. Three subspecies, H. elegans subsp. costaricensis Henderson, H. elegans subsp. sanblasensis Henderson, and H. elegans subsp. tacarcunensis Henderson are described as new, and two new combinations are made: H. elegans subsp. sodiroi (Dammer ex Burret) Henderson and H. elegans subsp. concinna (H. E. Moore) Henderson. One subspecies occurring in the Amazon region is complex morphologically and is not resolved by the methods used here.     

A biogeographical analysis of terrestrial isopods (Isopoda, Oniscidea) from the central Aegean islands (Greece)

Abstract. Terrestrial isopods (Isopoda, Oniscidea) of forty-three central Aegean islands and islets are analysed from a biogeographical point of view. The oniscid fauna consists of sixty-nine species, thirty-two genera and fourteen families and is characterized by Aegean elements that have an eastern Mediterranean origin. Endemism is at the level of 20%, similar to that of oniscids from other Mediterranean archipelagoes. Analysis of species distribution through both idiographic and quantitative approaches (faunal similarity among islands using Jaccard's index, simple matching indices and the UPGMA clustering technique) resulted in the recognition of a major disjunction between Kykladesplus-Ikaria and eastern islands-plus-Astypalea and four noteworthy island groups: the eastern islands, Naxos-Ikaria, Amorgos, and all other Kyklades. Island similarities are generally consistent with the known palaeogeography of the area with the exception of Ikaria's placement with Naxos among the Kyklades. Also, the 'eastern' character of Astypalea is in conflict with evidence from other known taxa.     

A new polyploidScilla (Liliaceae) from the Cretan area (Greece)

Scilla talosii Tzanoudakis etKypriotakis from the islet of Dia is described as a new species. It is an autumn-flowering taxon related toScilla autumnalis L. andS. obtusifolia Poiret but distinct morphologically and cytologically from both these taxa.S. talosii is polyploid with 2n=ca. 150 chromosomes.     

Palaeoreas,Majoreas, and Stryfnotherium gen. nov. (Mammalia: Artiodactyla: Bovidae) from the Late Miocene of Greece

A new comparative study of the two crania, LGPUT RZO-03 and LGPUT RZO-68 attributed to Palaeoreas zouavei Bouvrain, 1980 from the Late Miocene of Axios Valley, Greece reveals great morphological and metric differences between them, which require the exclusion of LGPUT RZO-68 from this species and even from Palaeoreas Gaudry, 1861. This decision leaves unsupported many diagnostic features of P. zouavei applied by previous authors and along with recent data from Bulgaria put in doubt the validity of P. zouavei as a whole. As a result, the genus Palaeoreas is re-defined as monotypic and the cranial morphometric changes of P. lindermayeri (Wagner, 1848) are discussed on a temporal basis. An updated taxonomic analysis confirms the presence of P. lindermayeri in the Late Turolian mammal assemblage of Dytiko-1 (Axios Valley, Greece) but rejects its occurrence at Dytiko-2. Instead, a re-examination of old cranial specimens and a study of new ones suggest the presence in both sites of Majoreas Kostopoulos, 2004 a Late Miocene taxon previously mostly known from the eastern banks of the Aegean Sea. A reassessment of the LGPUT RZO-68 cranium, and its comparison with Palaeoreas and Criotherium Forsyth-Major, 1891, suggests that it deserves taxonomic distinction at genus and species levels: Stryfnotherium exophthalmon.