The carnivoran community from the Miocene of Sandelzhausen (Germany)

From the Bavarian Early/Middle Miocene (MN5) site Sandelzhausen, nine species of carnivoran mammals are identified including the hemicyonine ursid Hemicyon stehlini, the amphicyonids Amphicyon cf. major and Pseudarctos bavaricus, the mustelids Ischyrictis zibethoides and Martes cf. munki, the mephitid Proputorius pusillus, the viverrid Leptoplesictis cf. aurelianensis, the felid Pseudaelurus romieviensis, and finally the recently described barbourofelid Prosansanosmilus eggeri. With these taxa present, Sandelzhausen shows a carnivoran community typical, though deprived, for the Lower to Middle Miocene of Europe, but different from roughly contemporary Mediterranean faunas such as those from Çandir or Pa?alar in Turkey.     

The beaver (Mammalia,Castoridae) from the Miocene of Sandelzhausen (southern Germany)

The Early/Middle Miocene boundary locality Sandelzhausen (MN5, southern Germany) yielded three beaver teeth: one lower incisor fragment and two strongly worn upper premolars, a right and a left one. The latter are so similar in wear, height, size, and enamel pattern that they probably represent one individual. Although the teeth have been referred to Chalicomys jaegeri Kaup, 1832 (e.g., Hugueney 1999, Fahlbusch 2003), it seems more likely that they belong to Steneofiber depereti Mayet, 1908. However, the assignment of the present material to either one of these species is difficult, because in the advanced wear stage some diagnostic characteristics cannot be assessed. As to ecological implications for the locality Sandelzhausen the taxonomic identity of these two beaver species may not be important. Both are known from riverine, lacustrine or swamp sediments, and it can be assumed that they usually lived in close vicinity to water. Thus, the presence of beaver remains supports the presence of water at Sandelzhausen. However, the teeth might have an allochthonous origin from creeks of the rising Alps.      

Devonian Gastropoda from the Dornap “Massenkalk” complex (Bergisches Land,Germany)

Seven Middle Devonian (latest Givetian or earliest Frasnian) gastropod species are described from the quarry of Oetelshofen near Dornap (Bergisches Land, northwestern Rheinisches Schiefergebirge, Germany). New species areMurchisonia dornapica, Araeonema osterholzensis, Plagiothyra isekei andMacrochilina drozdzewski. Additionally, a new subspecies,Euryzone delphinuloides oetelshofensis and aNaticopsis-like species and their typical facial occurrence are discussed. The existence of predation is proved by a bore-hole inMacrochilina drozdzewski; the predator itself is not yet identified with certainty.      

The Miocene <Emphasis Type="Italic">Keramidomys</Emphasis> (Rodentia,Eomyidae) from the Sandelzhausen locality (Germany)

In this note the taxonomic position of the tiny eomyid genus Keramidomys (Hartenberger, 1967) from the Early/Middle Miocene boundary locality of Sandelzhausen in the Bavarian Upper Freshwater Molasse is reexamined. As the chronological dating of the Sandelzhausen fossil site has been modified in the past from formerly the Neogene mammal unit MN6 to now MN5 and thanks to new abundant material this rodent is compared with other European forms. It is shown that the Sandelzhausen eomyid must be called K. thaleri Hugueney & Mein, 1968 on the basis of several morphological dental differences from K. carpathicus Schaub & Zapfe, 1953. This rodent seems to be an immigrant from East Asia into Europe. Even if K. thaleri is known in many European localities, all correlated to MN5, it is always a rare element of European rodent faunas. Differentiation from K. carpathicus is not easy and requires a sufficiently large sample.      

Continental molluscs of the fossil site Sandelzhausen (Miocene; Upper Freshwater Molasse from Bavaria) and their value for palaeoecological assessment

The continental mollusc fauna of Sandelzhausen (Southern Germany, Early/Middle Miocene, MN5) is presented and shown to consist of 66 snail and 3 bivalve taxa. Previous ecological interpretations were partly based on poorly preserved material and are shown to be erroneous. Within a series of quantitative samples, ecologically four mollusc communities can be distinguished (two each for terrestrial and freshwater molluscs), highly indicative of ecological conditions and changes in the landscape surrounding the fossil site as well as the locality itself. Open scrub-lands with temporary waters, as indicated by forms such as Granaria and the dominance of lymnaeids, are steadily replaced with damp forests with abundant litter and rotting wood surrounding a perennial lake with small tributaries. These more favorable conditions are indicated by an overall increase of mollusc diversity, rising abundance of planorbids, and presence of restricted forms such as Drepanotrema (intolerant to droughts) as well as certain forest dwellers such as Gastrocopta and hygromiids. Molluscs are highly sensitive to environmental changes, which is reflected in the fossil record of Sandelzhausen.      

Bats (Chiroptera, Mammalia) from Middle Miocene karstic fissure fillings of Petersbuch near Eichstätt, Southern Franconian Alb (Bavaria)

The bats from the karstic fissure fill sites Petersbuch 6, 10, 18, 31, 35 and 48 include 14 species, four of which are new. All samples are numerically dominated by rhinolophids, especially by Rhinolophus delphinensis. The rich samples from Petersbuch 6-18 are striking in their high species diversity (seven species each). Myotis bavaricus nov. sp. is characterised by a series of plesiomorphous characters, while Myotis reductus nov. sp. is more derived. A special trait of Submyotodon petersbuchensis nov. gen. nov. sp. is the coexistence of nyctalodont and (sub-) myotodont molars in one dentary. Miniopterus rummeli nov. sp. is characterised by its large size, a high coronoid process and its widely spaced lower premolars. Composition and origin of the chiropteran faunas are discussed. They are interpreted as an accumulation of bats that perished in the cave or rock crevice and to a negligible extent as remains of owl pellets. The stratigraphic correlation of the faunas, based on rodents, is late Middle Miocene (MN 7/8). This study is based on several thousand specimens.     

A partial skeleton of Prodeinotherium bavaricum (Proboscidea, Mammalia) from the Middle Miocene of Unterzolling (Upper Freshwater Molasse, Germany)

Here, a partial skeleton of Prodeinotherium bavaricum from Unterzolling (Southern Germany) is documented. The following elements are preserved and described for the first time: cervical vertebrae 1-2 and 5-7, the first thoracic vertebra, one lumbar vertebra, trapezium, metacarpals 1-5, tibia, calcaneus, endo- and mesocuneiform, cuboid, the fourth metatarsal, and some phalanges. Comparisons with the skeletons of P. bavaricum from Franzensbad (Czech Republic) and Deinotherium giganteum from Eserovo (Bulgaria) show osteological differences that are described and discussed.     

The diet of <Emphasis Type="Italic">Metaschizotherium bavaricum</Emphasis> (Chalicotheriidae,Mammalia) from the MN 5 of Sandelzhausen (Germany) implied by the mesowear method

The genus Metaschizotherium is a Miocene member of the Schizotheriinae (Chalicotheriidae, Perissodactyla) and appears in the MN 5 of Southern Germany with the species M. bavaricum. The Chalicotheriidae have mostly been reconstructed as browsers. In this study, the mesowear method is applied to 11 upper premolars and molars of M. bavaricum from the Upper Freshwater Molasse locality of Sandelzhausen (MN 5). With this method the amount of abrasive and attritive dental wear is investigated and thus it provides a time-averaged signature of food abrasiveness of ungulates. Principal components analysis is performed on mesowear variables of M. bavaricum. This species is found to classify closest to extant mixed feeding ruminants. This indicates that the diet of M. bavaricum essentially included nonabrasive browse but also a certain amount of abrasive plant material. The composition of the diet of extant reference species indicates that this abrasiveness was most likely imposed by bark and whole branches. In the palaeoecological context of Sandelzhausen M. bavaricum occupied the dietary niche of a mixed feeder or an abrasion-dominated browser. A high degree of similarity is recognised between the mesowear patterns of M. bavaricum from Sandelzhausen and M. fraasi from the MN 6 of the Franconian Alb. This is interpreted as reflecting a similar dietary niche of the two species.      

Studies on the effect of NAD(H) on nitrogenase activity in Rhodospirillum rubrum

The effect of NAD(P) and analogs of this nucleotide on nitrogenase activity in Rhodospirillum rubrum has been studied. Addition of NAD⁺ to nitrogen fixing Rsp. rubrum leads to inhibition of nitrogenase. NADP⁺ has the same effect but NADH or analogs modified in the nicotinamide portion do not cause inhibition. In contrast to ammonium ions, addition of NAD⁺ leads to inhibition of nitrogenase in cells that have been N-starved under argon. The inhibitory effect of NAD⁺ is more pronounced at lower light intensities. Addition of NAD⁺ also leads to inhibition of glutamine synthetase, a phenomenon also occurring when “switchoff” is produced by the addition of effectors such as ammonium ions or glutamine. It is also shown that NAD⁺ is taken up by Rsp. rubrum cells.     

Stable isotope ecology of Miocene large mammals from Sandelzhausen,southern Germany

The carbon, oxygen, and strontium isotope composition of enamel from teeth of large Miocene herbivorous mammals from Sandelzhausen (MN5, late Early/early Middle Miocene) in the North Alpine foreland basin, were analyzed to infer diet and habitat. The mean enamel δ¹³C value of −11.4 ± 1.0‰ (n = 53) for the nine taxa analyzed (including proboscideans, cervids, suids, chalicotheres, equids, rhinocerotids) indicates a pure C₃ plant diet for all mammals. ⁸⁷Sr/⁸⁶Sr ratios of ~0.710 higher than those from teeth of the western Molasse Basin (0.708–0.709) seem to indicate preferential feeding of the mammals in the northeastern Molasse Basin. The sympatric herbivores have different mean δ¹³C and δ¹⁸O values which support diet partitioning and/or use of different habitats within a C₃ plant ecosystem. Especially the three sympatric rhinoceroses Plesiaceratherium fahlbuschi, Lartetotherium sansaniense, and Prosantorhinus germanicus show clear partitioning of plants and/or habitats. The palaeomerycid Germanomeryx fahlbuschi was a canopy folivore in moderately closed environments whereas Metaschizotherium bavaricum (Chalicotheriidae) and P. germanicus (Rhinocerotidae) were browsers in more closed forest environments. The horse Anchitherium aurelianense was probably a more generalized feeder than assumed from its dental morphology. The forest hog Hyotherium soemmeringi has the highest δ¹³C and lowest δ¹⁸O value of all analyzed taxa, possibly related to a frugivorous diet. Most taxa were water-dependent browsers that record meteoric water δ¹⁸O values of about −5.6 ± 0.7‰ Vienna Standard Mean Ocean Water (VSMOW). Using a modern-day mean annual air temperature (MAT)– relation a MAT of 19.3 ± 1.5°C can be reconstructed for Sandelzhausen. A Gomphotherium subtapiroideum tusk serially sampled for δ¹⁸O values does not record a clear pattern of seasonality. Thus most taxa were C₃ browsers in a forested and humid floodplain environment in the Molasse Basin, which experienced a warm-temperate to subtropical climate and possibly low seasonality.      

Restudy and reassignment ofDentalium antiquum Goldfuss, 1841 (Middle Devonian)

Dentalium antiquum is one of two moderately well-known Devonian “scaphopods” in the German literature. Examination of the type material and a few specimens in other institutions indicates more individual variability than is to be expected in species of the molluscan class Scaphopoda. The species is transferred with question toColeolus Hall, a fossil presumed to be a calcareous “worm” tube.      

Merychippus (Mammalia, Perissodactyla, Equidae) from the Middle Miocene of state of Oaxaca, southeastern Mexico

Mexican material referable to Merychippus from two localities in eastern Oaxaca was described first nearly 50 years ago. Subsequent work there and in Central Oaxaca, spanning some 30 years, has allowed to establish the detail stratigraphy in both regions, and assembled a collection of merychippine material from the Matatlán (Central Oaxaca) and El Camarón (eastern Oaxaca) Formations, both K-Ar dated ~15 Ma (late early Barstovian). Detailed taxonomic analysis of this collection indicate the presence of two subhypsodont horse species referable to “Merychippus” cf. “M.” primus and “M.” cf. “M.” sejunctus in both regions. These records document the coexistence in tropical southern North America of basal and hipparionine affinity merychippine grade species, and provide a glimpse in to the diversity of subhypsodont equids in this region.     

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.     

Early Tithonian Saturnalidae (Radiolaria) from the Solnhofen area (Southern Franconian Alb, southern Germany)

In order to complete the study of the very rich early Tithonian (Hybonoticeras hybonotum Zone) radiolarian fauna from the Mühlheim Member of the Mörnsheim Formation outcropping in the Solnhofen area, the taxa of the family Saturnalidae are described. Although rather rare, the Saturnalidae of this member contain 14 species, ten of which are new. These species belong to four genera, one of which is new (Moebicircus n. gen.), and two subfamilies (Hexasaturnalinae and Saturnalinae). The taxonomy at generic level of these late Jurassic radiolarians is founded on the basis of the position of the blades along the ring and number and morphology of the spines. Type of spines (simple or forked) has either species level value or none, depending on species. Special attention was given to anomalies, which sometimes are rather frequent, since they can give Information of paleobiological and paleoecological Orders. Among them frequent cases of open ring and additional spines withDicerosaturnalis and Siamese twins skeletons withSpongosaturninus andDicerosaturnalis are to be noted. The authors hope that this new taxonomy will give a better image of the evolution and radiation of the Saturnalidae during the Tithonian.     

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.     

“Meganthropus” cranial fossils from Java

There are now twelve significant hominid cranial fossils from the Lower and Middle Pleistocene of Java, all but two being from the Sangiran site. Most of this material is well-known in the literature, but three skulls, possibly representing “Meganthropus” are here described in detail for the first time. Most scholars have assigned them all toHomo erectus, while others have suggested that they represent as many as four different hominoid taxa. The author argues that they represent two possible species of hominids. “Meganthropus” I, II, and III are more massive than any of the knownH. erectus specimens. They are also relatively higher vaulted, apparently smaller brained, and have unusually thick lower occipital planes. “Meganthropus” may represent a species that separated fromH. erectus upon its arrival to Java.     

Tsuga shimokawaensis,a new species of permineralized conifer leaves from the Middle Miocene Shimokawa Group,Hokkaido, Japan

Silicified pinaceous leaves from the late Middle Miocene bed of Shimokawa Town, central Hokkaido, Japan are described asTsuga shimokawaensis sp. nov. Comparisons with leaves of extant species ofTsuga show that the new species is assigned to sect.Tsuga, resembling the extant speciesT. heterophylla in having usually one-cell-layered hypodermis, andT. sieboldii in having mesophyll cells between the resin canal and the hypodermis. From cladistic analyses we infer thatTsuga shimokawaensis is positioned the basal in sect.Tsuga and that sect.Tsuga comprises four species groups.     

Albertona balkanica from Early Miocene of Snegotin (Serbia) – implications for the distribution and evolution of Miocene Ochotonidae

Remains of Albertona balkanica from the Early Miocene clays of eastern Serbia have shifted the range boundary of this species in direction of Central Europe. The peculiarities in tooth morphology were used for comparison and defining of the possible evolutionary connection with other representatives of fossil Ochotonidae. Due to the similarity with fauna of Aliveri in Greece, the association of small mammals of Snegotin (which includes Albertona) was included in the MN4 zone.