A DNA barcode library for Germany′s mayflies,stoneflies and caddisflies (Ephemeroptera,Plecoptera and Trichoptera)

Mayflies, stoneflies and caddisflies (Ephemeroptera, Plecoptera and Trichoptera) are prominent representatives of aquatic macroinvertebrates, commonly used as indicator organisms for water quality and ecosystem assessments. However, unambiguous morphological identification of EPT species, especially their immature life stages, is a challenging, yet fundamental task. A comprehensive DNA barcode library based upon taxonomically well‐curated specimens is needed to overcome the problematic identification. Once available, this library will support the implementation of fast, cost‐efficient and reliable DNA‐based identifications and assessments of ecological status. This study represents a major step towards a DNA barcode reference library as it covers for two‐thirds of Germany's EPT species including 2,613 individuals belonging to 363 identified species. As such, it provides coverage for 38 of 44 families (86%) and practically all major bioindicator species. DNA barcode compliant sequences (≥500 bp) were recovered from 98.74% of the analysed specimens. Whereas most species (325, i.e., 89.53%) were unambiguously assigned to a single Barcode Index Number (BIN) by its COI sequence, 38 species (18 Ephemeroptera, nine Plecoptera and 11 Trichoptera) were assigned to a total of 89 BINs. Most of these additional BINs formed nearest neighbour clusters, reflecting the discrimination of geographical subclades of a currently recognized species. BIN sharing was uncommon, involving only two species pairs of Ephemeroptera. Interestingly, both maximum pairwise and nearest neighbour distances were substantially higher for Ephemeroptera compared to Plecoptera and Trichoptera, possibly indicating older speciation events, stronger positive selection or faster rate of molecular evolution.     

Diagenesis of growth bands in fossil scleractinian corals: identification and modes of preservation

Light and dark bands in fully recrystallized fossil hermatypic corals are generally interpreted to represent annual growth increments reflecting a photosymbiotic life style—an interpretation of far reaching significance in palaeoecology. In this paper we describe annual growth bands in the colonial coral Porites in a perfect (aragonite and microstructures retained) and fully recrystallized (sparry calcite mosaic) style of preservation from sediments of Late Miocene age (Crete, Greece). Analysis of a continuous spectrum of transitional preservational stages shows that in Miocene Porites preservation of the growth banding was controlled by preferential dissolution of the high-density band associated with cementation by drusy calcite spar during freshwater diagenesis/shallow burial diagenesis. Marine precipitates (pelletoidal Mg-calcite) preferentially accumulated along tabulate dissepiments producing an additional growth rhythmicity. Massive Porites had annual growth rates of ∼4.0 mm, whereas in ramose branching Porites, a conspicuous banding is formed by concentrations of marine micropelletoidal cement along dissepiments at ∼1.8 mm spacing. If taken as annual growth increments, these bands represent very low extension rates, however, they may rather reflect subannual forcing functions (i.e., lunar cycles). An identical scenario of precipitation and concentration of pelletoidal carbonate along dissepiments and dissolution-controlled documentation of growth bands can be inferred for Late Jurassic microsolenids. Therefore, growth bandings in fossil corals potentially reflect both, monthly and annual cycles. Consequently, care must be taken when using coral growth bands in palaeoecology and palaeoclimatology.     

The Cenomanian of northern Germany: facies analysis of a transgressive biosedimentary system

A facies analysis of the epicontinental marine Cenomanian sediments of northern Germany shows the presence of 17 facies types (FTs, including several subtypes) which can be assigned to three facies associations: 1) an inner shelf facies association (FT 1–8) with high amounts of terrigenous material and/or high-energy depositional features, 2) a middle shelf facies association (FT 9–15) of predominantly calcareous sediments with moderate amounts of generally fine siliciclastics, and 3) an outer shelf facies association (FT 16–17) of low-energy, fine-grained, pure limestones. These three facies associations roughly correspond to the well-known lithological units of the Cenomanian of northern Germany, i.e., the Essen Greensand/Cenomanian Marls complex, the Pläner Limestones, and the Poor rhotomagense Limestones. The sediments were deposited on a northward-dipping homoclinal ramp with more-or-less shoreline-parallel facies belts. The sediment composition on this ramp-like shelf was a function of the varying importance of three different sediment sources: 1) terrigenous input from the south (Rhenobohemia), generally fining/decreasing in a proximal–distal (i.e., S–N) direction; 2) production of skeletal grains, mainly by macrobenthic organisms; and 3) settling of planktic carbonate (mainly calcispheres and calcareous nannofossils). In response to decreasing water energy with increasing water depth, the seaward decreasing terrigenous influence, and increasing planktic carbonate production, increasingly finer and more calcareous sediments were deposited in a proximal–distal transect. This rather straightforward picture was slightly modified by highest carbonate accumulation rates (planktic and benthic) on the middle shelf, forming a mid-shelf depocenter (fossiliferous, calcisphere-rich Pläner Limestones). Time-transgressive, southward-directed onlap of this biosedimentary system during the Cenomanian caused a significant retreat of the coastline towards the south and a retrogradational stacking of facies belts, explaining the broadly similar facies development and lithology of Cenomanian successions across northern Germany. The boundaries of the lithological units, however, tend to be considerably diachronous in a distal–proximal transect. In the late Middle and early Late Cenomanian, a final drowning and facies levelling (“oceanization”) is indicated by the widespread deposition of uniform calcareous nannofossil mudstones (Poor rhotomagense Limestones).     

Management of calcareous grasslands for Nickerl’s fritillary (<Emphasis Type="Italic">Melitaea aurelia</Emphasis>) has to consider habitat requirements of the immature stages,isolation, and patch area

We analysed the habitat preferences of adult stages and oviposition electivity of Melitaea aurelia in calcareous grasslands in the Diemel Valley (central Germany) to assess the key factors for successful management. Egg-laying and adult habitats of M. aurelia were more or less congruent. Oviposition electivity at the host plant (Plantago media) was best explained by a combination of host plant quantity and vegetation structure. Habitat quality, isolation and patch area explained 86% of the current patch occupancy of M. aurelia. With M. aurelia preferentially inhabiting transitional vegetation types, management requires a balance between abandonment and disturbance. Disturbances provide open soil that facilitates germination of the host plant Plantago media. On the other hand, immature and adult stages of M. aurelia perform best on calcareous grasslands with a high amount of host plants and low disturbance intensity. Traditional rough grazing regimes seem to be the most favourable tool for developing the necessary spatial and temporal heterogeneity in patches. The best results may be achieved by rotational grazing where only a subset of inhabited patches is grazed intensively each year. Our analysis of patch occupancy indicates that it would be desirable to restore patches in close proximity to occupied sites.     

Early jurassic (Upper Pliensbachian) gastropods from the Herforder Liasmulde (Bielefeld, Northwest Germany)

An Early Jurassic (Upper Pliensbachian) gastropod fauna from the Herforder Liasmulde (Herford Lias Syncline) near Bielefeld, Northwest Germany is described. This is one of the first detailed and illustrated reports of Pliensbachian gastropods from this area. Sixteen species are reported. Surface collecting yielded relatively large vetigastropods (pleurotomarioids amd trochomorphs). Bulk samples yielded abundant small caenogastropods (especiallyLevipleura blainvillei andKalchreuthia frankei) and cylindrobullinid opisthobranchs (e.g.Cylindrobullina dornend). These small species are also abundant in the Pliensbachian of South Germany (N Bavaria, Franconia). Therefore, this characteristic Pliensbachian microgastropod assemblage was widespread in Germany and probably also in other comparable marine deposits of Central Europe (e.g. the Paris Basin).      

Revision ofSemionotus bergeri Agassiz, 1833 (Upper Triassic, Germany), with comments on the taxonomic status ofSemionotus (Actinopterygii, Semionotiformes).

Semionotus bergeri Agassiz, 1833, from the Upper Triassic (Carnian) of southern Germany, the type species ofSemionotus, is here described in detail for the first time, including so far unstudied material in the Naturkunde-Museum Coburg, which allows a fairly complète reconstruction of this fish. The species is diagnosed by the présence of long frontals, about 4 times longer than wide, with antorbital lateral processes, 6–7 posterior infraorbitals, infraorbital at the posteroventral corner of the orbit notably larger than adjacent ones, infraorbitals at the posterior border of the orbit being as deep as long, and the middle élément anteroventrally expanded, dorsal fin base about two times the anal fin base.S. elegans from the Late Triassic-Early Jurassic of the Newark Supergroup in North America, is so far the best know species ofSemionotus and is the only species of this genus that has been included in phylogentic analyses. However, the comparison ofS. bergeri with other proposed speciesof Semionotus, such asS. elegans, and other semionotids casts doubt on the monophyly of this genus as currently understood, since these species exhibit a mosaic distribution of characters.      

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.     

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.      

Upper Pleistocene Panthera leo spelaea (Goldfuss, 1810) remains from the Bilstein Caves (Sauerland Karst) and contribution to the steppe lion taphonomy,palaeobiology and sexual dimorphism

Remains of the steppe lion Panthera leo spelaea (Goldfuss) from historical digs in the Bilstein Caves of Warstein (Sauerland, NW Germany) are described. Their age seems to be from the Early Weichselian periods (Upper Pleistocene). Whereas the Bilstein cave was inhabited by cave bears at that time only a few hyena prey remains, were most likely imported into the cave entrance by hyenas. Bite and crush marks on a few bones of Bison priscus, Bos primigenius, Cervus elaphus, a rhinoceros Coelodonta antiquitatis vertebra and even several chewed cave bear bones prove the hyena presence which is similar to other caves in the Sauerland hyena den cave rich region. Additionally some larger wolves subspecies Canis lupusspelaeus bones were found, but only few Crocuta crocuta spelaea remains are present. After taphonomic comparisons to six other hyena and cave bear den caves of northern Germany, this cave can be classified as a cave bear den, which was briefly used by hyenas only for food storage or commuting or cave bear predation site in one part of the Cave. The lion material refers at least to one young adult lioness, one more adult female and two male lions; therefore, at minimum, the remains of four adult individuals are represented. The absence of juvenile lion material, in contrast to cave bear cub remains in the Bilstein Caves, proves that P. leo spelaea did not use this and all other caves in the region to raise their cubs. The bone material from the Bilstein Caves would prove the same hyena-lion antagonism conflict being recently proven for the Perick Caves, Balve Cave or Martins Cave well. Other situations in caves such as the Keppler Cave and the Bilstein Cave initially show the more complex taphonomic situation of lion remains in European caves, especially in cave bear dens, where they seem to have hunted periodically cave bears, such as it is already proven for hyenas in the Sauerland Karst and other caves of Europe.