A Late Miocene Paratethyan mollusc fauna from the Denizli Basin (southwestern Anatolia, Turkey) and its regional palaeobiogeographic implications

We report on a Late Miocene mollusc fauna from a single locality in the Denizli Basin in southwestern Turkey that is composed of ten presumably euryhaline species and three freshwater species. The fauna is remarkably distinct from faunas of the adjacent Late Miocene Aegean and Euxinian Basins, and has elements in common with the modern Caspian Sea fauna in the form of Didacna species. The suggested age of the fauna (Late Miocene) would extend the stratigraphic range of the lymnocardiid genus Didacna (hitherto Calabrian-Extant) considerably. The palaeobiogeographic significance of the Denizli fauna is discussed.     

Miocene honey bees from the Randeck Maar of southwestern Germany (Hymenoptera, Apidae)

The Miocene Randeck Maar (southwestern Germany) is one of the only sites with abundant material of fossil honey bees. The fauna has been the focus of much scrutiny by early authors who recognized multiple species or subspecies within the fauna. The history of work on the Randeck Maar is briefly reviewed and these fossils placed into context with other Tertiary and living species of the genus Apis Linnaeus (Apinae: Apini). Previously unrecorded specimens from Randeck Maar were compared with earlier series in an attempt to evaluate the observed variation. A morphometric analysis of forewing venation angles across representative Recent and Tertiary species of Apis as well as various non-Apini controls was undertaken to evaluate the distribution of variation in fossil honey bees. The resulting dendrogram shows considerable variation concerning the wing venation of Miocene Apini, but intergradation of other morphological characters reveals no clear pattern of separate species. This suggests that a single, highly variable species was present in Europe during the Miocene. The pattern also supports the notion that the multiple species and subspecies proposed by earlier authors for the Randeck Maar honey bee fauna are not valid, and all are accordingly recognized as Apis armbrusteri Zeuner.     

Stratigraphic interpretation of the Kulu Formation (Early Miocene, Rusinga Island, Kenya) and its implications for primate evolution

Early Miocene fossils from Rusinga Island, Kenya, provide some of the best evidence for catarrhine evolution and diversification, and, together with more than eighty-five other mammalian species, form an important comparative reference for understanding faunal succession in East Africa. While there is consensus over the stratigraphic position of most of Rusinga's volcaniclastic deposits, the lacustrine Kulu Formation has been placed in various parts of the geological sequence by different researchers. To resolve this discrepancy, we conducted detailed geological analyses which indicate that the Kulu Formation was formed in the Early Miocene during a period of volcanic inactivity and subsidence following the early, mainly explosive hyper-alkaline phase of the Kisingiri complex and prior to the final eruptions of nephelinitic lavas. The underlying Hiwegi and older formations were locally deformed and deeply eroded before sedimentation began in the Kulu basin, so that the Kulu sediments may be significantly younger than the 17.8 Ma Hiwegi Formation and not much older than the overlying Kiangata Agglomerata-Lunene Lava series, loosely dated to ca. 15 Ma. The overall similarities between Kulu and Hiwegi faunas imply long-term ecological stability in this region. Our stratigraphic interpretation suggests that the Kulu fauna is contemporaneous with faunas from West Turkana, implying that differences between these assemblages—particularly in the primate communities—reflect paleobiogeographic and/or paleocological differences. Finally, the position of the Kulu Formation restricts the time frame during which the substantial faunal turnover seen in the differences between the primate and mammalian communities of Rusinga and Maboko Islands could have occurred.     

Paleoenvironmental reconstruction of middle Miocene paleosols bearing Kenyapithecus and Victoriapithecus, Nyakach Formation, southwestern Kenya

Paleosols in the middle Miocene (15 Ma) Nyakach Formation at Kaimogool, near Sondu, southwestern Kenya have yielded specimens of the early cercopithecoid Victoriapithecus macinnesi and the early kenyapithecine Kenyapithecus africanus, and can be used as evidence for the environmental mosaic occupied by these primates. Five distinct types of paleosols (pedotypes) are recognized in the Nyakach Formation section at Kaimogool South. The most common paleosols are reddish brown, silty calcareous profiles with blocky structure and large root traces (Ratong pedotype) which are interpreted as soils of well-drained, dry bushland or thicket (nyika). Weakly developed paleosols associated with paleochannels (Dhero pedotype) represent wooded grassland early in the ecological succession from streamside flooding. One of these paleosols has yielded a fossil flora of grasses and small-leaved dicots like those of modern semi-arid wooded grassland. Crumb structured, calcareous paleosols with iron-manganese nodules (Yom pedotype) are interpreted to represent seasonally waterlogged, wooded grassland (dambo or vlei). Thick, red clayey, calcareous paleosols with blocky ped structure and large root traces (Tut pedotype) are interpreted as soils of well-drained dry woodland. Other blocky-structured, gray to brown calcareous paleosols with iron-manganese nodules (Chido pedotype) are interpreted as soils of seasonally waterlogged, riparian dry woodland. Fossil soils, plants and gastropods are evidence of an unusually dry (300-500 mm mean annual precipitation) habitat for apes, consisting of a vegetational mosaic dominated by dry woodland, bushland and thickets with few areas of seasonally waterlogged grassland. Fossils of V. macinnesi are rare from Nyakach, but were found in paleosols representative of bushland and thicket habitats (Ratong). Fossils of the ape K. africanus were found within paleosols indicative of dry woodland (Tut). Other paleosol types representative of seasonally dry dambo grassland (Yom), colonizing grassland (Dhero) or riparian woodland (Chido) are also represented, but have not yet produced primate fossils.     

First hominoid from the Late Miocene of the Irrawaddy Formation (Myanmar)

For over a century, a Neogene fossil mammal fauna has been known in the Irrawaddy Formation in central Myanmar. Unfortunately, the lack of accurately located fossiliferous sites and the absence of hominoid fossils have impeded paleontological studies. Here we describe the first hominoid found in Myanmar together with a Hipparion (s.l.) associated mammal fauna from Irrawaddy Formation deposits dated between 10.4 and 8.8 Ma by biochronology and magnetostratigraphy. This hominoid documents a new species of Khoratpithecus, increasing thereby the Miocene diversity of southern Asian hominoids. The composition of the associated fauna as well as stable isotope data on Hipparion (s.l.) indicate that it inhabited an evergreen forest in a C3-plant environment. Our results enlighten that late Miocene hominoids were more regionally diversified than other large mammals, pointing towards regionally-bounded evolution of the representatives of this group in Southeast Asia. The Irrawaddy Formation, with its extensive outcrops and long temporal range, has a great potential for improving our knowledge of hominoid evolution in Asia.     

Early Miocene cricetids (Rodentia) from the Junggar basin (Xinjiang, China) and their biochronological implications

Among the 14 small mammal species from the early Miocene locality of the northern area of the Junggar basin (northern Xinjiang, China), four species are cricetids (Rodentia): unnamed species of Cricetodon and Eumyarion, and two new species, Karydomys debruijni nov. sp. and Megacricetodon beijiangensis nov. sp. Some aspects of the morphology of Cricetodon sp. are shared by Eucricetodon from the late Oligocene, suggesting that these specimens could be of intermediate form between Eucricetodon and Cricetodon. One tooth of Eumyarion sp. was found, making its determination uncertain, but its morphology is clearly differentiated from the one of Cricetodon sp. The species K. debruijni nov. sp. is established based on its primitive features compared to the species known in Europe and Anatolia, and its specific association of characters compared to Karydomys dzerzhinskii. M. beijiangensis nov. sp. shows many plesiomorphic features compared to the species already described in the middle Miocene of China. Based on both the whole assemblage of rodents and the species of cricetids, the biochronologic position and the age of the locality are discussed. The locality appears to be biochronologically very close to the fauna from the Chul’adyr Formation in Aktau Mountains, but we propose an age slightly older than the one proposed for this fauna, probably equivalent to the MN3 biozone in Europe.     

Effects of fragments and landscape characteristics on the orchid bee richness (Apidae: Euglossini) in an urban matrix,southwestern Amazonia

Orchid bees are important pollinators in tropical forests. Although studies have already detected effects of habitat loss and forest fragmentation on bee assemblages, little is known about orchid bees in urban forest fragments. The aim of this study was to analyse the influence of forest fragments (size and edge index) and landscape features (forest cover area and built-up area around the forest fragments, connectivity and spatial distance from the urban center) on the abundance, richness and composition of orchid bees. Male bees were attracted by odoriferous baits and collected in ten forest fragments of different sizes. In total, we collected 3166 male bees belonging to 4 genera and 38 species. The increase of the built-up area and the reduction of the forest cover area around the forest fragments decreased the abundance and richness of bees. We recorded a smaller number of bees in areas closer to the urban center. We failed to find a significant relationship between abundance/richness of bees and forest fragment area, edge index, and connectivity. We observed that the faunistic dissimilarity was positively correlated with the geographic distance between forest fragments. The forest fragments that presented greater landscape dissimilarity also presented greater faunistic dissimilarity. Our results suggest that the matrix structure around the forest fragments is an important factor that influences the Euglossini bee assemblages inside these forest fragments. Based on our results, we believed that the conservation of fragments with a larger forest cover area and smaller built-up areas around them contribute to the maintenance of the diversity of orchid bees and their pollination services.     

The Structure of the Mammalian Predator Guild in the Santa Cruz Formation (Late Early Miocene)

The Santa Cruz Formation (late early Miocene, Santacrucian age) registers 11 species of mammalian predators (Metatheria, Sparassodonta). Together with large carnivorous flightless birds, they comprised the terrestrial predator guild. The Santacrucian sparassodonts were diverse in body size, had different locomotory habits, and were primarily hypercarnivores. The objective of this work is to analyze the guild structure of the sparassodonts of the Santa Cruz Formation, using the variables of body mass, diet, and locomotion as proxies. Furthermore, we analyze the interaction with other predators and potential prey. The univariated test V of Poole and Rathcke and the multivariated test of Clark-Evans were used to construct the models. In the multivariate test, we made a Principal Component Analysis to resume and standardize the variables. With body mass and locomotion we obtained an evenly spaced pattern of segregation for the sparassodont species, being non-significant and significant, respectively. The pattern was aggregated and significant only with diet. The analysis of all variables together resulted in an evenly spaced and significant pattern, which is consistent with character displacements (segregation of species throughout the morphospace) that would help to diminish interspecific competition during the Santacrucian age and would allow selection of prey species of different sizes and substrate specializations. When the body size pattern of predator birds and sparassodonts were plotted together, the pattern is evenly spaced and non-significant. Other factors, including locomotion, would differentiate these species and their ecological niches.     

哈萨克斯坦中部(Kaydagul Formation) 欧洲水松的角质层结构分析

研究了哈萨克斯坦中部上渐新统/下中新统的欧洲水松(Glyptostrobus europaeus (Brongn.) Ung.)的枝叶微形态结构,并利用扫描电子显微镜和光学显微镜与"现存最近亲缘种"--水松(G. pensilis C. Koch)的表皮特征进行了比较.     

Palaeoecology of the Miocene El Cien Formation (Mexico) as determined from wood anatomical characters

In order to establish the general environmental conditions under which the plants of the El Cien Formation grew during the Miocene, wood anatomical features from several extant ecological categories (southern California flora) and communities (Brazil, Mexico and Suriname) as well as those of the El Cien paleocommunity were analysed using different statistical approaches. We find a significant correlation between 42 characters of 199 species and 11 climate variables which suggests a close association of wood characters with temperature, whereas characters such as diameter, frequency, and grouping of vessels are more closely related to evaporation. The expression of various wood characters depends on more than one climate variable. There are no doubts regarding the tropical affiliation of the El Cien fossil flora but water availability and seasonality of the system are less clear. Principal component and cluster analyses of two different datasets (423 species, 9 characters and 223 species, 32 characters) suggest that the physiognomy of the fossil assemblage of El Cien was highly efficient in water transport, but also highly vulnerable to embolisms. Characters such as vessel diameter, frequency and grouping suggest proximity to a tropical rain forest, whereas the proportion of ring semi ring porous woods brings the El Cien paleoflora closer to forests with slightly xeromorphic physiognomy, probably a tropical deciduous or semi-deciduous forest. This condition is paralleled with the taxonomic composition of the El Cien fossil plants since their Nearest Living Relatives (NLR) can be referred to both humid or more (seasonally) dry environments. Derived regression analysis models suggest that the paleoflora prospered under conditions with an evaporation regime similar to that of the tropical rain forest.     

Geometric morphometric analysis of a new Miocene bumble bee from the Randeck Maar of southwestern Germany (Hymenoptera: Apidae)

The first fossil bumble bee (Apinae: Bombini) from the Miocene Randeck Maar of southwestern Germany is described and illustrated. The specimen is subjected to a geometric morphometric analysis along with a diversity of other bumble bee species representing most major extant lineages, and particularly the subgenus Bombus s.s. The morphometric analysis supports the placement of the Randeck Maar species within Bombus s.s., as a species distinct from all others in the subgenus. It shows that extant subgenera of bumblebees were already derived in the early/middle Miocene. The Randeck Maar fossil is formally described as Bombus (Bombus) randeckensis Wappler & Engel sp. n. .     

Miocene Sanitheriidae (Suiformes, Mammalia) from Namibia and Kenya: systematic and phylogenetic implications

Sanitheres are enigmatic small suoids with bunoselenodont cheek teeth and a tendency for complication of the premolars by polycuspy and polycristy. Until a few years ago virtually nothing was known about their anterior dentition, but recent discoveries in Greece, Kenya and Namibia have thrown light on their incisor and canine morphology, and reveals among other things that the canines are highly sexually dimorphic, and the dm/1 is replaced by a p/1. Material from Kipsaraman, Kenya, collected between 1997 and 2003, consists of parts of mandibles and many isolated teeth, including elements of the deciduous dentition, a significant proportion of which are unworn. These new finds confirm the aberrant nature of the sanithere dentition within a suoid framework, and support their classification as a family separate from Suidae and Palaeochoeridae (= Old World Tayassuidae).

Résumé

Les sanithères sont de petits Suoïdes aux dents jugales bunosélénodontes avec des prémolaires qui présentent une tendance à la multiplication des cuspides et des crêtes (“polycuspidie” et la “polycristie”). Jusqu’à récemment, on ne connaissait rien sur la morphologie de leur dentition antérieure, mais des découvertes récentes réalisées en Grèce, au Kenya et en Namibie, lèvent le voile sur la morphologie de leurs incisives et de leur canine. Ainsi, il apparaît que les canines sont très dimorphes et que la dm/1 inférieure est remplacée par une p/1. Le matériel de Kipsaraman au Kenya, récolté entre 1997 et 2003, consiste en des fragments de mandibules et de nombreuses dents isolées, dont des dents déciduales, qui pour la plus grande part ne sont pas usées. Ces nouveaux fossiles confirment la nature aberrante de la dentition des sanithères au sein des Suoidea, et supporte leur classification dans une famille distincte des Suidae, et des Palaeochoeridae (= Tayassuidae de l’Ancien Monde).     

An additional example of the ichnogenus Zoophycos in Cambrian strata (Gallatin Formation; Wyoming,USA)

The ichnogenus Zoophycos is relatively rare in Cambrian-age sedimentary rocks, being far more commonly encountered in post-Devonian, and particularly post-Paleozoic, strata. A new occurrence of Zoophycos from the Upper Cambrian Gallatin Formation of the Bighorn Mountains, north-central Wyoming, USA, is reported herein. Material consists of a single specimen preserved within quartz silt representing the upper surface of a graded quartz arenite bed. The burrow is a relatively wide, curved, horizontally oriented lobe with a poorly visible marginal tube but well-developed internal lamellae. Discovery of this specimen contributes to the paleobiogeographic distribution of lower Paleozoic Zoophycos, adds to the literature suggesting that most Cambrian Zoophycos were horizontal and shallow-tier structures, and provides further support to the notion that relatively complex deposit-feeding behavior evolved earlier in the Phanerozoic than had long been recognized.     

The Cyatheaceae and Dicksoniaceae (Pteridophyta) of Bolivia

A summary is presented of all Bolivian species of Cyatheaceae and Dicksoniaceae. In total two species of Dicksoniaceae (Culcita 1 sp.,Dicksonia 1 sp.) and 34 species of Cyatheaceae (Sphaeropteris 1 sp.,Alsophila 5 spp.,Cyathea 26 spp.,Cnemidaria 2 spp.) are known. One hybrid inCyathea is recognized. The endemicCyathea dintelmannii is newly described, andCyathea herzogii from Bolivia and Peru is separated fromCyathea caracasana var.boliviana. Both species are illustrated. An artificial key to the Bolivian species of Cyatheaceae is provided.     

Miocene bristlemouths (Teleostei: Stomiiformes: Gonostomatidae) from the Makrilia Formation,Ierapetra, Crete

Bristlemouths of the genus Cyclothone are currently regarded as the most abundant vertebrates on Earth. The fossil record seems to suggest that these fishes diversified during the Miocene in the Pacific Ocean, but there is no evidence of their presence in the Miocene of the Atlantic Ocean and Mediterranean basin. A new bristlemouth, Cyclothone gaudanti sp. nov. (Teleostei, Stomiiformes, Gonostomatidae), is described herein based on 16 specimens from the Upper Miocene Makrilia Formation (late Tortonian of Crete, Greece). The small sized species is characterized by light pigmentation, 30–31 (14–15 + 15–16) vertebrae, dorsal fin with 10–13 rays, anal fin with 10–14 rays, premaxilla bearing seven closely spaced teeth, maxilla with 42–55 teeth, epipleurals, and autogenous parhypural. The presence of epipleurals appears to be unique of this Miocene species, and the re-establishment of this ancestral character state may be possibly interpreted as related to a phylogenetic character reversal. Morphological and paleoecological considerations suggest that this species possibly inhabited the upper mesopelagic layer, at depths ranging from 2–300 and 500 meters.