Origin and Early Evolution of Hydrocharitaceae and the Ancestral Role of Stratiotes.

The combined morphological features of Stratiotes (Hydrocharitaceae) pollen, observed with light and electron microscopy, make it unique among all angiosperm pollen types and easy to identify. Unfortunately, the plant is (and most likely was) insect-pollinated and produces relatively few pollen grains per flower, contributing to its apparent absence in the paleopalynological record. Here, we present fossil Stratiotes pollen from the Eocene of Germany (Europe) and Kenya (Africa), representing the first reliable pre-Pleistocene pollen records of this genus worldwide and the only fossils of this family discovered so far in Africa. The fossil Stratiotes pollen grains are described and compared to pollen from a single modern species, Stratiotes aloides L. The paleophytogeographic significance and paleoecological aspects of these findings are discussed in relation to the Hydrocharitaceae fossil records and molecular phylogeny, as well as the present-day distribution patterns of its modern genera.

Plastid DNA sequences and oospore characters of some European taxa of Tolypella section Tolypella (Characeae) identify five clusters, including one new cryptic Tolypella taxon from Sardinia, but they do not coincide with current morphological descriptions.

In Europe, the genus Tolypella (Characeae) comprises four to eight Tolypella taxa in sections Rothia and Tolypella that have been distinguished by vegetative morphology and gametangial characters such as antheridial size and oospore wall ornamentation. However, morphological differentiation is difficult in some cases due to overlapping and variable vegetative features, which in many cases are difficult to observe clearly. To clarify the taxonomic status of the five European taxa of Tolypella in section Tolypella, sequence data of the plastid genes atpB, rbcL and psbC for Tolypella glomerata (Desv.) Leonh., Tolypella hispanica Allen, Tolypella nidifica (O.F. Müll.) A. Braun, Tolypella normaniana (Nordst.) Nordst. and Tolypella salina Cor. were combined with data on oospore morphology, including oospore wall ornamentation. Gene sequence data identified five distinct clusters, but they were not consistent with the morphologically identified five taxa. T. glomerata consisted of some of the samples morphologically identified as T. glomerata and seven samples of T. normaniana, while the remaining T. glomerata samples clustered with specimens of unclear affiliation (Tolypella sp.). We identified two clusters of T. hispanica within the European material: cluster T. hispanica I consisted of samples from various locations, whereas the second cluster (T. hispanica II) consisted of samples of T. hispanica from Sardinia Island. The remaining cluster consisted of all the specimens that had been determined as T. salina or T. nidifica in addition to two specimens of T. normaniana. Oospore morphology was most clearly distinguishable for T. glomerata. Oospore characteristics for all other taxa were not as informative but showed some geographical and/or environmentally influenced differences, especially for T. nidifica and T. salina. Our results suggest the need to further check the different taxonomy of Tolypella sect. Tolypella in which specimens normally identified as T. glomerata might be two different taxa, T. glomerata and an unidentified taxon; T. nidifica and T. salina are not separate taxa; T. normaniana is a diminutive variant of two different Tolypella taxa; and T. hispanica comprises two different taxa, one from the Mediterranean island Sardinia.

Greenhouse conditions in lower Eocene coastal wetlands?-Lessons from Schöningen, Northern Germany.

The Paleogene succession of the Helmstedt Lignite Mining District in Northern Germany includes coastal peat mire records from the latest Paleocene to the middle Eocene at the southern edge of the Proto-North Sea. Therefore, it covers the different long- and short-term climate perturbations of the Paleogene greenhouse. 56 samples from three individual sections of a lower Eocene seam in the record capture the typical succession of the vegetation in a coastal wetland during a period that was not affected by climate perturbation. This allows facies-dependent vegetational changes to be distinguished from those that were climate induced. Cluster analyses and NMDS of well-preserved palynomorph assemblages reveal four successional stages in the vegetation during peat accumulation: (1) a coastal vegetation, (2) an initial mire, (3) a transitional mire, and (4) a terminal mire. Biodiversity measures show that plant diversity decreased significantly in the successive stages. The highly diverse vegetation at the coast and in the adjacent initial mire was replaced by low diversity communities adapted to wet acidic environments and nutrient deficiency. The palynomorph assemblages are dominated by elements such as Alnus (Betulaceae) or Sphagnum (Sphagnaceae). Typical tropical elements which are characteristic for the middle Eocene part of the succession are missing. This indicates that a more warm-temperate climate prevailed in northwestern Germany during the early lower Eocene.

Eocene fossil is earliest evidence of flower-visiting by birds.

Birds are important pollinators, but the evolutionary history of ornithophily (bird pollination) is poorly known. Here, we report a skeleton of the avian taxon Pumiliornis from the middle Eocene of Messel in Germany with preserved stomach contents containing numerous pollen grains of an eudicotyledonous angiosperm. The skeletal morphology of Pumiliornis is in agreement with this bird having been a, presumably nectarivorous, flower-visitor. It represents the earliest and first direct fossil evidence of flower-visiting by birds and indicates a minimum age of 47 million years for the origin of bird-flower interactions. As Pumiliornis does not belong to any of the modern groups of flower-visiting birds, the origin of ornithophily in some angiosperm lineages may have predated that of their extant avian pollinators.

Testing for the effects and consequences of mid paleogene climate change on insect herbivory.

BACKGROUND: The Eocene, a time of fluctuating environmental change and biome evolution, was generally driven by exceptionally warm temperatures. The Messel (47.8 Ma) and Eckfeld (44.3 Ma) deposits offer a rare opportunity to take a census of two, deep-time ecosystems occurring during a greenhouse system. An understanding of the long-term consequences of extreme warming and cooling events during this interval, particularly on angiosperms and insects that dominate terrestrial biodiversity, can provide insights into the biotic consequences of current global climatic warming. METHODOLOGY/PRINCIPAL FINDINGS: We compare insect-feeding damage within two middle Eocene fossil floras, Messel and Eckfeld, in Germany. From these small lake deposits, we studied 16,082 angiosperm leaves and scored each specimen for the presence or absence of 89 distinctive and diagnosable insect damage types (DTs), each of which was allocated to a major functional feeding group, including four varieties of external foliage feeding, piercing- and-sucking, leaf mining, galling, seed predation, and oviposition. Methods used for treatment of presence-absence data included general linear models and standard univariate, bivariate and multivariate statistical techniques. CONCLUSIONS/SIGNIFICANCE: Our results show an unexpectedly high diversity and level of insect feeding than comparable, penecontemporaneous floras from North and South America. In addition, we found a higher level of herbivory on evergreen, rather than deciduous taxa at Messel. This pattern is explained by a ca. 2.5-fold increase in atmospheric CO(2) that overwhelmed evergreen antiherbivore defenses, subsequently lessened during the more ameliorated levels of Eckfeld times. These patterns reveal important, previously undocumented features of plant-host and insect-herbivore diversification during the European mid Eocene.

Aromatized arborane/fernane hydrocarbons as biomarkers for cordaites.

Previous palaeobotanical and palynological studies on coals from Euramerican Pennsylvanian ( identical with Late Carboniferous) coal basins indicate a major change in coal-swamp floras, especially at the Westphalian-Stephanian ( approximately Kasimovian-Gzhelian, according to Geological Time Scale 2004) boundary. A flora dominated by arborescent lycophytes was replaced by a vegetation dominated by marattialean tree ferns in various Euramerican coal basins. Earlier combined palynological and organic geochemical studies on Westphalian/Stephanian coals and shales from the Saar-Nahe Basin (Germany) revealed that the distribution of aromatized arborane/fernane hydrocarbons in solvent extracts reflects the increasing importance of seed plants, especially cordaites (extinct group of gymnosperms), conifers and pteridosperms. However, the biological source of the precursor molecules could not be specified. To clarify if the arborane/fernane derivatives MATH, MAPH, DAPH 1, and DAPH 2 in Westphalian/Stephanian coals can be assigned to one of the three potential source plant groups, we analyzed coals, sediments and fossil plant remains from different Euramerican locations with respect to their biomarker composition and stable carbon isotopic composition. Thereby, stable carbon isotopic ratios showed only insignificant variations between Westphalian and Stephanian samples and proved to be an unsuitable tool to describe floral changes during the Westphalian/Stephanian of the Saar-Nahe Basin. In contrast, we were able to show for the first time that MATH, MAPH, DAPH 1 and DAPH 2 are prominent constituents only in extracts of cordaitean macrofossils and can therefore be regarded as biomarkers for this group of gymnosperms.