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Earlier studies indicate a strong correlation of pollen morphology and ultrastructure with taxonomy in Loranthaceae. Using high-resolution light microscopy and scanning electron microscopy imaging of the same pollen grains, we document pollen types of 35 genera including 15 studied for the first time. Using a molecular phylogenetic framework based on currently available sequence data with good genus-coverage, we reconstruct trends in the evolution of Loranthaceae pollen and pinpoint traits of high diagnostic value, partly confirming earlier intuitive hypotheses based on morphological observations. We find that pollen morphology in Loranthaceae is strongly linked to phylogenetic relationships. Some pollen types are diagnostic for discrete genera or evolutionary lineages, opening the avenue to recruit dispersed fossil pollen as age constraints for dated phylogenies and as independent data for testing biogeographic scenarios; so far based exclusively on modern-day data. Correspondences and discrepancies between palynological and molecular data and current taxonomic/systematic concepts are identified and suggestions made for future palynological and molecular investigations of Loranthaceae.

Dating the subsidence history of the North Atlantic Land Bridge (NALB) is crucial for understanding intercontinental disjunctions of northern temperate trees. Traditionally, the NALB has been assumed to have functioned as a corridor for plant migration only during the early Cenozoic, but recent fi ndings of plant fossils and inferences from molecular studies are challenging this view. Here, we report dispersed pollen of Quercus from Late Miocene sediments in Iceland that shows affi nities with extant northern hemispheric white oaks and North American red oaks. Older (15 to 10 Ma) sediments do not contain pollen of Quercus suggesting it arrived after that time. Pollen from the 9 -8 Ma Hr ú tagil locality is indistinguishable from morphotypes common among white and red oaks. In contrast, pollen from the 5.5 Ma Sel á rgil locality has a tectum that is at present confi ned to North American white and red oaks, indicating a second episode of migration to Iceland. These fi ndings suggest that transatlantic migration of temperate plant taxa may not have been limited by vast areas of sea or by cold climates during the Miocene. Furthermore, our results offer a plausible explanation for the remarkably low degree of genetic differentiation between modern disjunct European and North American oaks.

In this paper we document Fagaceae pollen from the Eocene of western Greenland. The pollen record suggests a remarkable diversity of the family in the early Cenozoic of Greenland. Extinct Fagaceae pollen types include Eotrigonobalanus, which extends at least back to the Paleocene, and two ancestral pollen types with affinities to the Eurasian Quercus Group Ilex and the western North American Quercus Group Protobalanus. In addition, modern lineages of Fagaceae are unambiguously represented by pollen of Fagus, Quercus Group Lobatae/Quercus, and three Castaneoideae pollen types. These findings corroborate earlier findings from Axel Heiberg Island that Fagaceae were a dominant element at high latitudes during the early Cenozoic. Comparison with coeval or older mid-latitude records of modern lineages of Fagaceae shows that modern lineages found in western Greenland and Axel Heiberg likely originated at lower latitudes. Further examples comprise (possibly) Acer, Aesculus, Alnus, Ulmus, and others. Thus, before fossils belonging to modern northern temperate lineages will have been recovered from older (early Eocene, Paleocene) strata from high latitudes, Engler’s hypothesis of an Arctic origin of the modern temperate woody flora of Eurasia, termed ‘Arcto-Tertiary Element’, and later modification by R. W. Chaney and H. D. Mai (‘Arcto-Tertiary Geoflora’) needs to be modified.

The fossil record provides good evidence for the minimum ages of important events in the diversification and geographic spread of Asteridae, with earliest examples extending back to the Turonian stage of the Late Cretaceous (~89 Ma). Some of the fossil identifications accepted in previous considerations of asterid phylogeny do not stand up to careful scrutiny. Nevertheless, among major clades of asterids, there is good evidence for a range of useful anchor points. Here we provide a synopsis of fossil occurrences that we consider as reliable representatives of modern Asterid families and genera. In addition, we provide new examples documented by fossil dispersed pollen investigated by both light and scanning electron microscopy studies including representatives of Loranthaceae, Amaranthaceae, Cornaceae (incl. Nyssa L., Mastixia Blume, Diplopanax Hand.-Mazz.), Sapotaceae, Ebenaceae, Ericaceae, Icacinaceae, Oleaceae, Asteraceae, Araliaceae, Adoxaceae and Caprifoliaceae from Paleogene sites in Greenland, western North America, and central Europe, and of Lamiaceae and Asteraceae from the middle to late Miocene northeastern China. We emphasize that dispersed pollen, taken along with megafossil and mesofossil data, continue to fill gaps in our knowledge of the paleobotanical record.

ABSTRACT. Modern lineages of the beech family, Fagaceae, one of the most important north-temperate families of woody flowering plants, have been traced back to the early Eocene. In contrast, molecular differentiation patterns indicate that the Fagus lineage, Fagoideae, with a single modern genus, evolved much earlier than the remaining lineages within Fagaceae (Trigonobalanoideae, Castaneoideae, Quercoideae). The minimum age for this primary split in the Fagaceae has been estimated as 80 ± 20 Ma (i.e. Late Cretaceous) in recently published, time-calibrated phylogenetic trees including all Fagales. Here, we report fagaceous fossils from the Campanian of Wyoming (82-81 Ma; Eagle Formation [Fm]), the Danian of western Greenland (64-62 Ma; Agatdal Fm), and the middle Eocene of British Columbia (ca 48 Ma; Princeton Chert), and compare them to the Fagaceae diversity of the recently studied middle Eocene Hareøen Fm of western Greenland (42-40 Ma). The studied assemblages confirm that the Fagus lineage (= Fagoideae) and the remainder of modern Fagaceae were diverged by the middle Late Cretaceous, together with the extinct Fagaceae lineage(s) of Eotrigonobalanus and the newly recognised genus Paraquercus, a unique pollen morph with similarities to both Eotrigonobalanus and Quercus. The new records push back the origin of (modern) Fagus by 10 Ma and that of the earliest Fagoideae by 30 Ma. The earliest Fagoideae pollen from the Campanian of North America differs from its single modern genus Fagus by its markedly thicker pollen wall, a feature also seen in fossil and extant Castaneoideae. This suggests that a thick type 1 foot layer is also the plesiomorphic feature in Fagoideae although not seen in any of its living representatives. The Danian Fagus pollen of Greenland differs in size from those of modern species but is highly similar to that of the western North American early Eocene F. langevinii, the oldest known beech so far. Together with the Quercus pollen record, absent in the Campanian and Danian formations but represented by several types by the middle Eocene, this confirms recent dating estimates focussing on the genera Fagus and Quercus, while rejecting estimates from all-Fagales-dated trees as too young. The basic Castaneoideae pollen type, still found in species of all five extant genera of this putatively paraphyletic subfamily, represents the ancestral pollen type of most (modern) Fagaceae (Trigonobalanoideae, Castaneoideae, Quercoideae).

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Shallowing of the Panama