Rudabànya: Taphonomic analysis of a fossil hominid site from Hungary

The late Miocene deposits at Rudabànya, Hungary, were laid down in a shallow valley sloping westwards from a range of hills and opening out into the Pannonian Lake. Rise and fall of lake level gave rise to varying conditions, from dry land with soil formation to swamp and lake. The stratigraphic and palaeontological succession has been investigated at one of the sites, Rudabànya 2, where two cycles of deposition and erosion are represented, with soil formation, swamp conditions with lignite formation, and periods of extended high lake level succeeding each other. Both mammal and plant fossils are present at several levels. Taphonomic modifications in the Rudabànya 2 vertebrate faunas include losses of skeletal elements through carnivore selection, fluvial sorting at some levels, and post-depositional destruction by leaching and/or acid soils. The lowest level, the lower lignite, has few fossils. The fossil mammals from the level above, the grey marl, are the least modified but they are mixed with abraded, probably allochthonous, bone fragments and more complete specimens resulting from near-lake deaths. Modifications of bones by carnivores are indicated, but the specimens were too broken post-depositionally for the impact of the carnivores to be assessed. Carnivore action is also indicated for the fauna of the black clay which formed on the surface of the grey marl. The fauna consists of relatively abundant small mammals and the primates Anapithecus hernyaki and Dryopithecus hungaricus, with the latter much less common. The predator accumulating the smaller species was probably a viverrid. The red marl fauna is a transported assemblage from higher up the valley with the fossils extremely fragmentary and abraded and few identifiable specimens, almost all of which are teeth. The black mud fauna is also probably a transported assemblage, lower energy than the red marl environment, and the bones are much modified subsequently by acid corrosion similar to that seen today in bone preserved in peat bogs. Dryopithecus is a major constituent of the fauna, with Anapithecus less common. D. hungaricus is thus associated more strongly with swamp forest and shallow riverine conditions with low energy movement of water, and A. hernyaki is associated with lake shore (probably forest) conditions, accumulating in lake sediments and lake-flat sediments. The palaeoecology of the area as a whole, based on the associated flora and fauna, is a combination of swamp forest, lake shore forest and open mud flats.     

New evidence for diet and niche partitioning in Rudapithecus and Anapithecus from Rudabánya,Hungary

Rudabánya is rare among Eurasian Miocene fossil primate localities in preserving both a hominid and pliopithecoid, and as such provides the unique opportunity to reconstruct the nature of sympatry and niche partitioning in these taxa. Rudapithecus and Anapithecus have similar locomotor and positional behavior and overlapping body mass ranges. While prior analyses of molar occlusal anatomy and microwear identify Rudapithecus as a soft-object frugivore, reconstructing the dietary behavior of Anapithecus has been more problematic. This taxon has been interpreted to be more folivorous by some, and more frugivorous by others. Here, we use high-resolution polynomial curve fitting (HR-PCF) to quantify and evaluate the mesiodistal and cervico-incisal curvatures of the incisor crowns of Rudapithecus and Anapithecus to identify diet-specific morphological variation in these taxa. Results are consistent with the interpretation that Anapithecus and Rudapithecus were primarily frugivorous and had diets that included similar resource types. However, Anapithecus may have consumed greater amounts of foliage, similar to extant mixed folivore–frugivores (i.e., Gorilla gorilla gorilla, Symphalangus syndactylus), while Rudapithecus generated elevated compressive loads in the incisor region consistent with a specialized role for the anterior dentition in food processing (i.e., removal of tough protective fruit pericarps). We interpret these findings in light of the paleoecology at Rudabánya and conclude that, if these taxa were indeed sympatric, Anapithecus may have used additional leaf consumption as a seasonal fallback resource to avoid direct competition with Rudapithecus. Conversely, Rudapithecus may have relied on less preferred and harder fruiting resources as a seasonal fallback resource during periods of fruit scarcity.     

Taxonomic and functional implications of stigma morphology in species ofCassia,Chamaecrista, andSenna (Leguminosae:Caesalpinioideae)

Two stigma forms occur inChamaecrista andSenna, but only one inCassia. In the common chambered form, a stigma pore is positioned on the reflexed style tip and is the entrance to a tapering chamber. The pore rim is fringed by hairs which vary in number, size, distribution and shape. In the alternative form the stigma is situated at the apex of the curved style and is crateriform. The crater rim is fringed by hairs of variable number and shape. The stigmatic hairs are predominantly unicellular and cutinized. Stigma and hair differences aid in the taxonomy of the genera. Their functions in pollination biology are discussed.     

Isolation ofNosema slovaca (Microsporidiae) fromDermacentor reticulatus ticks (Acari: Ixodidae) collected in Hungary

Nosema slovaca (Microsporidiae) was isolated from a maleDermacentor reticulatus tick collected in the Mélykút locality, Bács-Kiskun province, Hungary. This parasitic protozoan caused an acute infection in partly engorgedD. reticulatus females when inoculated intracoelomically resulting in death of the ticks within 5–15 days post-infection. This strain ofNosema can be maintained in the laboratory by passaging between partly engorgedD. reticulatus females and is currently being studied for its potential as a biological control agent of ixodid ticks.     

Fossil frogs (Anura) from Shanwang (Middle Miocene; Shandong Province, China)

The early middle Miocene biota from the Shanwang locality in eastern China is one of the most diverse and well preserved in the world. This overall richness contrasts with its low diversity of anuran amphibians. The most abundant anurans are representatives of the family Ranidae, which remains the dominant group in the extant anuran fauna of China. Besides Rana basaltica Young, some other ranids can be recognised at the locality, and are clearly distinguishable from R. basaltica based on their larger size and different proportions. Ranids are also represented by a developmental series of tadpoles, ranging from approximately Nieuwkoop-Faber (NF) stage 43 to a metamorphosing tadpole of NF stage 57. This is the first known developmental series of fossil non-pipoid Anura outside the Pelobatidae. In addition to the ranids, Bufo shandongensis nov. sp. (replacement name for B. linquensis), represented by one of the earliest known articulated bufonid skeletons, is described and illustrated. Another large toad, Macropelobates linquensis (Yang) nov. comb., is re-described; it can be assigned to the Pelobatoidea, with a possible relationship to the Scaphiopodidae, and can be considered, based on the presence of a large spade, a tibiofibula shorter than the femur, and a short astragalus and calcaneus, to be the earliest known anuran burrower.     

Chromosome numbers and taxonomic implications in the fern genusAzolla (Azollaceae)

Investigation of chromosome numbers of allAzolla species, and for the first time of hybrids, has been undertaken. Removal of wax from the leaf surface proved invaluable in achieving clear cytological preparations and providing unambiguous chromosome numbers. In contrast to previous records, the speciesA. pinnata, A. filiculoides, A. filiculoides var.rubra, A. caroliniana, A. microphylla, andA. mexicana were found to be 2n=44, andA. nilotica to be 2n=52. Several triploids (2n=66) and one tetraploid (2n=88) were identified. No geographical pattern could be observed in the distribution of triploids which probably derive from the function of unreduced gametes. The chromosome number of hybrids occasionally deviates from the diploid chromosome number (2n=44). The small chromosome size limits karyotypic analysis and only differences in overall chromosome size can be observed. Taxonomic implications of chromosome numbers and sizes are discussed.     

A new hominoid hamate and first metacarpal from the Late Miocene Nagri Formation of Pakistan

A hamate and the proximal part of a first metacarpal from the type locality of the Nagri Formation in Pakistan, and attributed to Sivapithecus parvada, are described. In overall proportions, the hamate is rather robust, showing most similarity to that of Gorilla. Unlike extant hominoids it lacks a well-developed hamulus, and its triquetral facet is morphologically dissimilar to that in extant anthropoids. The morphology of the hamate indicates effective weight transmission through the ulnar side of the wrist, limited ulnar deviation and restricted extension in the triquetrohamate joint, and stability of the hamatometacarpal joints. The morphology of the partial first metacarpal is most similar to that of Pan. Previously described postcranial bones of S. parvada indicate that its locomotor behaviour included both quadrupedalism and climbing. This is consistent with the limited evidence of the first metacarpal, whereas the hamate strongly emphasizes the quadrupedal aspect of the locomotor repertoire.     

Dentition ofSivaladapis nagrii (Adapidae) from the late Miocene of India

Two genera and three species of adapid primates are known from the middle and late Miocene of India and Pakistan. Most fossil specimens are fragmentary, but the best-known species, Sivaladapis nagrii,is now represented by enough specimens to permit composite reconstruction of much of the dentition. The incisors of Sivaladapishave spatulate crowns, and the canines are large, projecting teeth. Premolars and molars exhibit complex occlusion involving simultaneous approximation of pointed leading cusps on upper and lower molars, with linear trailing lophs. The premolar eruption sequence in Sivaladapisappears to be P ₂-P₄-P₃, as in most extant prosimians. Symphyseal fusion of the mandibular rami occurred early in ontogeny, before the eruption of any of the anterior permanent teeth. We interpret Sivaladapisto have been a specialized arboreal folivore that became extinct near the end of the Miocene, when the distribution of forests was increasingly restricted and colobine monkeys first invaded South Asia.     

Taxonomic implications of Liassic ferns Cladophlebis Brongniart and Todites Seward from Hungary

Ferns are diverse and abundant in the Lower Liassic flora of the Mecsek Mountains in south Hungary. The morphogenus Cladophlebis Brongniart, 1849, and the genus Todites Seward, 1900, show a high diversity. In the examined material, most of the characteristics used in taxonomy vary, showing transitional forms that cast doubt on the separation of species established on the grounds of either a low number of specimens, a low number of features or uncharacteristic differences.     

Taxonomic and phytogeographic implications from ITS phylogeny in<Emphasis Type="Italic"> Berberis</Emphasis> (Berberidaceae)

A phylogeny based on the internal transcribed spacer (ITS) sequences from 79 taxa representing much of the diversity of Berberis L. (four major groups and 22 sections) was constructed for the first time. The phylogeny was basically congruent with the previous classification schemes at higher taxonomic levels, such as groups and subgroups. A notable exception is the non-monophyly of the group Occidentales of compound-leaved Berberis (previously separated as Mahonia). At lower levels, however, most of previous sections and subsections were not evident especially in simple-leaved Berberis. Possible relationship between section Horridae (group Occidentales) and the simple-leaved Berberis clade implies paraphyly of the compound-leaved Berberis. A well-known South America-Old World (mainly Asia) disjunctive distribution pattern of the simple-leaved Berberis is explained by a vicariance event occurring in the Cretaceous period. The ITS phylogeny also suggests that a possible connection between the Asian and South American groups through the North American species (Berberis canadensis or B. fendleri) is highly unlikely.     

Lagomorphs from the Miocene of Sandelzhausen (southern Germany)

Three genera of lagomorphs, Prolagus, Lagopsis, and “Amphilagus,” were identified during a revision of the lagomorph material from Sandelzhausen (MN5, Early/Middle Miocene boundary, southern Germany). Evidence of two morphological and dimensional classes were observed at some tooth positions in Prolagus (some p3 show an unmistakable P. oeningensis morphology, others closely resemble P. crusafonti), but not at other tooth positions (e.g., M1–2). Insufficient data from Sandelzhausen precludes identification of two different species of Prolagus from this locality, and to define the characteristics of the possible P. crusafonti-like species. Thus, all Prolagus specimens have been classified as P. aff. oeningensis. The genus Lagopsis is represented by L. cf. penai, whose presence is compatible with a MN5 age. The relative abundance of Lagopsis to Prolagus may indicate relatively cool and wet palaeoclimatic conditions. The largest primitive lagomorph species from continental Europe is present at Sandelzhausen. Morphological and dimensional comparisons with other European primitive lagomorphs exclude any affinity with the genera Eurolagus and Titanomys and with the species included in “Amphilagus ulmensis”. Some common features with “Amphilagus antiquus” were observed, although they are not sufficient for the attribution to this taxon. Until there is a general revision of European primitive lagomorphs, the Sandelzhausen giant lagomorph is classified as “Amphilagus” sp. Its origins, whether from evolution within Europe or migration from Asia, remain unknown.      

New hyalellids (Crustacea, Amphipoda, Hyalellidae) from Lake Titicaca

Two new species of hyalellid amphipods, Hyalella crawfordi and H. gauthieri, are described from Lake Titicaca; H. echinus (Faxon, 1876) is redescribed. The H. echinus group of species is newly proposed for these three species, and a group diagnosis is provided. A key to the three species in the group is provided Full article published online at http://www.senckenberg.de/odes/06-10.htm.     

First records of lice (Insecta: Phthiraptera) from wild southern pudu, <Emphasis Type="Italic">Pudu puda</Emphasis> (Mammalia: Artiodactyla: Cervidae)

The presence of two species of Phthiraptera, Bovicola caprae (Gurlt, 1843) (Ischnocera: Bovicoliidae) and Solenopotes binipilosus (Fahrenholz, 1916) (Anoplura: Linognathidae), is reported for the first time from Pudu puda (Molina, 1782).     

Flavonol pentaglycosides of Cordyla (Leguminosae: Papilionoideae: Swartzieae): distribution and taxonomic implications

A survey of foliar flavonoids in the swartzioid legume genus Cordyla s.l. revealed that three species, C. haraka, C. pinnata and C. richardii, were rich in flavonol pentaglycosides. Their structures were elucidated by spectroscopic and chemical methods as the 3-O-alpha-l-rhamnopyranosyl(1-->3)-alpha-l-rhamnopyranosyl(1-->2)[alpha-l-rhamnopyranosyl(1-->6)]-beta-d-galactopyranoside-7-O-alpha-l-rhamnopyranosides of quercetin and kaempferol (cordylasins A and B, respectively). These compounds were not found in the remaining species, C. africana, C. densiflora, C. madagascariensis (two subspecies) and C. somalensis, which exhibited different profiles of flavonoid glycosides. The distribution of flavonol pentaglycosides in Cordyla s.l. does not support a recent proposal to place both C. haraka and C. madagascariensis in the genus Dupuya [Kirkbride, J.H., 2005. Dupuya, a new genus of Malagasy legumes (Fabaceae). Novon 15, 305-314]. The generic relationship between Cordyla s.l. and Mildbraediodendron is also reassessed on the basis of chemical characters, as the O-linked tetrasaccharide that characterises cordylasins A and B is the same as that found in mildbraedin (kaempferol 3-O-alpha-l-rhamnopyranosyl(1-->3)-alpha-l-rhamnopyranosyl(1-->2)[alpha-l-rhamnopyranosyl(1-->6)]-beta-d-galactopyranoside), the main foliar flavonoid of Mildbraediodendron excelsum. Mildbraedin itself was found to be a minor constituent of leaflet extracts of C. haraka, C. pinnata and C. richardii, and a major constituent of C. somalensis.     

New species ofBeaufortia andChamaelaucium (Myrtaceae),Drosera (Droseraceae) andPultenaea (Fabaceae) from SW. Australia

Four new species of the generaBeaufortia, Chamaelaucium, Drosera, andPultenaea are described based on the author's collections from SW. Australia. Their affinities are discussed and full illustrations are provided.Dedicated to Hofrat Univ.-Prof.K. H. Rechinger on the occasion of his 80th birthday.     

New Permian tetrapod footprints and macroflora from Turkey (Çakraz Formation, northwestern Anatolia): Biostratigraphic and palaeoenvironmental implications

New tetrapod footprints belonging to the ichnogenus Hyloidichnus have been discovered in Turkey for the first time, in the lower part of the Çakraz Formation (Northwestern Anatolia) and together with macrofloral imprints of Annularia and Stigmaria. These discoveries confirm the Permian age of the fossiliferous red beds in which the coniferophyte Walchia was previously recorded. Based on the stratigraphic range of Annularia, Stigmaria and Hyloidichnus known elsewhere, a Cisuralian age is proposed for these beds. These new ichno- and macrofloral remains, together with the sedimentological data (mudcracks, rain drops) suggest the presence of captorhinid reptiles living in a palustrine floodplain environment, and under a warm, seasonal climate alternating between humid and relatively long dry seasons. These climatic conditions may have permitted the migration of these captorhinids through Laurasia during the Permian.     

The antennae of neopseustid moths: Morphology and phylogenetic implications, with special reference to the sensilla (Insecta, Lepidoptera, Neopseustidae)

The antennae of Lepidoptera Neopseustidae were examined with the scanning electron microscope. The studied species, Nematocentropus cfr. omeiensis, Neopseustis meyricki, Synempora andesae, Apoplania valdiviana and Apoplania penai possess nine types of antennal flagellum sensilla: multiporous large sensilla basiconica, multiporous thin sensilla basiconica, multiporous small sensilla basiconica, multiporous sensilla trichodea, multiporous sensilla coeloconica; uniporous sensilla chaetica; aporous sensilla chaetica, aporous stylus-shaped sensilla chaetica, aporous sensilla styloconica.The multiporous sensillum type here termed “multiporous large sensillum basiconicum” is unknown from other Lepidoptera and probably constitutes an autapomorphy of the family Neopseustidae. This sensillum type is remarkable by having a single base in female Apoplania and Synempora while in male Apoplania it has a bifid or trifid base, and in male Synempora it is composed of two or three incompletely separated hairs. This may be the first recorded example of a sexually dimorphic lepidopteran sensillum type. The stylus-shaped sensillum chaeticum is a primitive type which occurs only in some lower Lepidoptera.     

A new species of Anchitherium (Equidae: Anchitheriinae) from the Middle Miocene of Abocador de Can Mata (Vallès-Penedès Basin, NE Iberian Peninsula)

Anchitherium nievei sp. nov. is erected on the basis of dentognathic remains from the late Aragonian (MN7+8, ca. 12.3–12.1 to 11.7–11.6 Ma, Middle Miocene) of three localities from the Abocador de Can Mata local stratigraphic series (els Hostalets de Pierola, Vallès-Penedès Basin, Catalonia, Spain). This taxon differs from endemic species of Anchitherium from the inner Iberian basins, most closely resembling A. aurelianense, A. hippoides and A. steinheimense by its marked brachyodonty and small dental size, although differing by dental proportions and several details of occlusal morphology. From a paleoenvironmental viewpoint, the presence of this taxon fits well with previous inferences that indicated a relatively closed and humid forested habitat, more similar to that of similarly aged French localities.     

Secondary spore formation in <Emphasis Type="Italic">Orchesellaria mauguioi</Emphasis> (Asellariales,Trichomycetes) and its taxonomic and ecological implications

Orchesellaria (Orc.) mauguioi (Asellariales) was detected from the hindguts of Isotomurus palustris (Collembola, Hexapoda) collected in Japan. The secondary spore formation on the exuviae of its host is described. When the host molted, its exoskelton including the ectoderm-originated hindgut cuticle to which Orc. mauguioi mycelia attaches was shed. Arthrospores of Orc. mauguioi germinated and penetrated the molt skin and produced secondary spores terminally. The secondary spore is a narrowly ovoid monosporangious sporangium accompanied by a capitate-lageniform terminal cell; these fall off together as a dispersive unit. The terminal cell is sterile and bears one to three fine filamentous extensions seven times as long as the length of the cell. In contrast to the appendages produced endogenously by other harpellids, the filament is produced exogenously and is similar to those of the genus Orphella (Harpellales). The taxonomic and ecological implications of the secondary spore formation of Orc. mauguioi are discussed by comparison with those of other Trichomycetes.