Friðgeir Grímsson

Episodic migration of oaks to Iceland: Evidence for a North Atlantic “land bridge” in the latest Miocene

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 findings 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 affinities 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 Hrútagil locality is indistinguishable from morphotypes common among white and red oaks. In contrast, pollen from the 5.5 Ma Selárgil locality has a tectum that is at present confined to North American white and red oaks, indicating a second episode of migration to Iceland. These findings 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.

Pollen, fruits, and leaves of Tetracentron (Trochodendraceae) from the Cainozoic of Iceland and western North America and their palaeobiogeographic implications

Dispersed pollen, fruits, and leaves of Tetracentron (Trochodendraceae) are described from the Miocene of Iceland and assigned to a new species, Tetracentron atlanticum. The Icelandic fossils represent the first unambiguous record of Tetracentron for the Cainozoic of Europe. Well‐preserved dispersed grains of Tetracentron are also identified from the Eocene and Miocene of western North America and compared to the Icelandic fossils. In general, Tetracentron pollen is rather uniform through the Eocene to Recent, only displaying minor variation in ornamentation. Leaves are diagnostic at the species level. The findings add substantial new information to the generally poor fossil record of Tetracentron and indicate a more circumpolar distribution of the genus than previously assumed. The north‐eastern Asian – western North American Cainozoic distribution for plant taxa presently confined to East Asia is relatively common. In contrast, the extension to Iceland is remarkable, particularly in view of the absence of the genus in the fossil record from eastern North America, Greenland, and Central Asia and mainland Europe.

Fagaceae pollen from the early Cenozoic of West Greenland: revisiting Engler’s and Chaney’s Arcto-Tertiary hypotheses

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.

Assessing the Fossil Record of Asterids in the Context of Our Current Phylogenetic Framework1

Abstract 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 million years ago [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 reliable representatives of modern asterid families and genera. In addition, we provide new examples documented by fossil-dispersed pollen investigated by both LM and SEM studies including representatives of Loranthaceae, Amaranthaceae, Cornaceae (including 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 in 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.

Lythrum and Peplis from the Late Cretaceous and Cenozoic of North America and Eurasia: new evidence suggesting early diversification within the Lythraceae.

PREMISE OF THE STUDY To fully understand the evolution of todays angiosperms, the fossil record of plant families and genera must be used to determine their time of origin and phytogeographic history. As within many angiosperm families, the interrelationships of extant Lythraceae are hard to resolve without sufficient data from the geological past. Here we establish the earliest fossil occurrences of Lythraceae and start resolving the interrelationships and evolution of two of its genera, Lythrum and Peplis. METHODS We studied several Cretaceous and Cenozoic palynofloras from the northern and southern hemispheres. Using the single-grain technique, we screened the treated samples for Lythrum- and Peplis-type pollen. The same individual pollen grains were observed under both the light- and scanning electron microscope, allowing a high taxonomic resolution to be achieved. KEY RESULTS Fossil Lythraceae pollen grains are rare in palynological samples. Nevertheless, we were able to identify Lythrum and Peplis pollen from Late Cretaceous sediments and thereby extend the fossil record of the two genera by ca 70 million years. CONCLUSIONS The appearance of Lythrum and Peplis in North America and Peplis in Asia at approximately the same interval in the mid Late Cretaceous points to an already wide geographical distribution by then. These findings add vital information for the time of origin of the Lythraceae and suggest a higher diversity within the family. They also indicate that the distribution of particular genera during the Cretaceous was wider than previously thought.

Evolutionary trends and ecological differentiation in early Cenozoic Fagaceae of western North America

UNLABELLED • PREMISE OF THE STUDY The early Cenozoic was a key period of evolutionary radiation in Fagaceae. The common notion is that species thriving in the modern summer-dry climate of California originated in climates with ample summer rain during the Paleogene.• METHODS We investigated in situ and dispersed pollen of Fagaceae from the uppermost Eocene Florissant fossil beds, Colorado, United States, using a combined light and scanning electron microscopy approach.• KEY RESULTS Pollen types of Castaneoideae with affinities to modern Castanea, Lithocarpus, and Castanopsis were recognized. Pollen of the extinct genus Fagopsis represents a derived type of Castaneoideae pollen. Infrageneric groups of Quercus were well represented, including pollen of Group Protobalanus. The taxonomic diversity of Fagaceae and of the total plant assemblage indicates a mosaic of microclimates, that range from pronounced to weakly seasonal climates and depend on slope aspect and elevation. Continental climatic conditions may have triggered the evolution of sclerophyllous leaves and adaptive radiation in Quercus and other taxa thriving today under distinctly summer-dry and winter-dry climates.• CONCLUSIONS Vegetation types similar to modern vegetation belts of the Coastal Ranges (chaparral, nemoral conifer forest) were established in the Front Range in the late Eocene. Coeval plant assemblages from the Coastal Ranges of California indicate distinctly subtropical, moist climates. Hence, characteristic elements found today in the summer-dry and winter-dry climates of Pacific North America (Quercus Group Protobalanus, Notholithocarpus) may opportunistically have dispersed into their modern ranges later in the Cenozoic. This scenario is in contrast to the evolution and migration patterns of their western Eurasian Mediterranean counterparts (Quercus Group Ilex).

Morphological Trends in the Fossil Pollen of Decodon and the Paleobiogeographic History of the Genus

Various pollen morphotypes of Decodon from the Cenozoic of North America, Europe, and Asia are described for the first time, using a combination of LM and SEM. The report includes the earliest pollen record of Decodon from the Northern Hemisphere. The pollen grains were obtained from sedimentary samples using the single grain technique, allowing for a high taxonomic resolution. The fossil pollen show clear traceable changes in morphology toward the present. The apparent pollen diversity noted at the end of the Cenozoic indicates that during the Miocene, Decodon was not a monotypic genus, as at present. This is also apparent from the macrofossil record summarized here. The combined fossil record of the genus is used to evaluate the paleogeographic distribution of Decodon in time and place and to speculate about its migration across and between continents in the light of major geological events.

Specialized and Generalized Pollen-Collection Strategies in an Ancient Bee Lineage

Iconic examples of insect pollination have emphasized narrowly specialized pollinator mutualisms such as figs and fig wasps and yuccas and yucca moths. However, recent attention by pollination ecologists has focused on the broad spectra of pollinated plants by generalist pollinators such as bees. Bees have great impact for formulating hypotheses regarding specialization versus generalization in pollination mutualisms. We report the pollination biology of six northern European species of an extinct tribe of pollen-basket-bearing apine bees, Electrapini, of early-middle Eocene age, examined from two deposits of 48 and 44 million years in age. These bees exhibit a pattern of generalized, incidental pollen occurring randomly on their heads, thoraces, and abdomens, obtained from diverse, nectar-bearing plants. By contrast, a more restricted suite of pollen was acquired for metatibial pollen baskets (corbiculae) of the same bee taxa from a taxonomically much narrower suite of arborescent, evergreen hosts with uniform flower structure. The stereotyped plant sources of the specialist strategy of pollen collection consisted of pentamerous, radially symmetrical flowers with a conspicuous gynoecium surrounded by prominent nectar reward, organized in structurally similar compound inflorescences. Pollen specialization in bees occurs not for efficient pollination but rather in the corbiculate Electrapini as food for bee larvae (brood) and involves packing corbiculae with moistened pollen that rapidly loses viability with age. This specialist strategy was a well-developed preference by the early Eocene, providing a geochronologic midpoint assessment of bee pollen-collection strategies.

Diverse fossil Onagraceae pollen from a Miocene palynoflora of north-east China: early steps in resolving the phytogeographic history of the family

The origin and evolution of angiosperms can be unravelled by using fossil records to determine first occurrences and phytogeographic histories of plant families and genera. Many angiosperm families, for example the Onagraceae, have a poor macrofossil record, but are more common in palynological records. Modern Onagraceae produce pollen clearly distinct from that of other angiosperms. Combined morphological features obtained by use of light and scanning electron microscopy have enabled assignment of fossil Onagraceae pollen to extant genera, and therefore tracing of the origin and past distributions of extant Onagraceae lineages. We studied a Miocene palynoflora from the Daotaiqiao Formation of north-east China. Using the single-grain technique, we examined individual Onagraceae pollen/tetrads using both light and scanning electron microscopy. Fossil Onagraceae pollen is more frequent than macrofossil remains, but is still rare, and usually represented by a single taxon in palynological samples. Remarkably, samples from the Miocene of north-east China contain five different species: two of Circaea, one of Epilobium, and two of Ludwigia. Such a large number of Onagraceae taxa from a single palynoflora is unknown elsewhere. Whereas Ludwigia pollen is known from Cenozoic sediments of the northern hemisphere, the Circaea pollen is the first fossil pollen assignable to this extant genus. This is also the first fossil record of Epilobium from China. Although the young geological age of this sample does not enable consideration of time of origin for the genera encountered, the co-occurrence of Circaea, Epilobium, and Ludwigia in the mid to late-Miocene of East Asia sheds some light on their phytogeographic histories.

The Biogeographic History of Iceland – The North Atlantic Land Bridge Revisited

Plants lacking long distance dispersal mechanisms required a functioning land bridge to colonize Iceland, a route provided by the North Atlantic Land Bridge (NALB). During the Cainozoic, the NALB, also referred to as the Thulean route, came into existence in the latest Paleocene and Early Eocene, but there has been considerable debate about the timing of its termination. The North Atlantic Land Bridge consisted of the well defined subaerial Greenland-Scotland Transverse Ridge. The individual parts of this ridge may have undergone markedly different subsidence histories during the Neogene. At the western end of the NALB, possible links between Greenland and North America are provided by the shallow bathymetric sill at the Davis Strait between southern Baffin Land and southwestern Greenland, and, alternatively, the more northern land connection between the Queen Elizabeth Islands and Greenland. In this chapter, we use evidence from different disciplines (geology, palaeontology, phylogeography), amended with a large new palaeobotanical data set emerging from the present study, to evaluate the history of the North Atlantic Land Bridge and its potential role for transatlantic plant migration during the Neogene.