A Late Messinian Palynoflora with a Distinct Taphonomy more

Chapter 9 A Late Messinian Palynoflora with a Distinct Taphonomy Abstract A thin, white coloured ash/pumice layer on top of the poor macrofossil units at the Selárgil locality yields a rich late Messinian palynoflora that was deposited under a markedly different taphonomic setting than most other late Cainozoic floras discussed in this book. Pollen contained in this volcanic sediment apparently was deposited in a very short time during the actual ash fall, whereas in most other Cainozoic formations in Iceland pollen was deposited in clastic sediments over a longer time interval. The unusual taphonomy of the Selárgil pollen and spores assemblage possibly acted as filter against insect pollinated woody species such as Rhododendron subsect. Pontica and other tree taxa that would otherwise be expected (Juglandaceae, Ulmaceae). Overall, the arboreal flora at Selárgil contains a number of partly warmth-loving relict taxa from older formations (Cathaya, Scyadopitys, Tetracentron) and some newcomers (Quercus, two types of Ericaceae). The occurrence of a new type of Quercus with clear biogeographic affinities to North America points to a functioning land bridge between Iceland and Greenland during the Late Miocene and to climatic conditions in northern North America and Greenland that would have allowed for migration of oaks to Iceland. 9.1 Introduction The Fnjóskadalur Formation is ca 5.5 Ma in age (late Messinian, latest Late Miocene). The sediments exposed at the Selárgil locality were first described by Pjetursson (1905). Much later, Sigurðsson (1975) gave a more complete description of the strata, including volcanic constructions, sedimentary types and structures, and origin and formation of different units. Sigurðsson also collected plant macrofossils and figured them in his work; these fossils represent the only known collection from this locality. Interestingly, from these sediments, Sigurðsson (1975) reported the first and so far only findings of freshwater bivalves from the late Cainozoic of Iceland. Around the same time, Akhmetiev et al. (1978) conducted a more regional investigation of the area and traced the occurrence of the Fnjóskadalur Formation (sediments as well as volcanic constructions). They also studied the T. Denk et al., Late Cainozoic Floras of Iceland, Topics in Geobiology 35, DOI 10.1007/978-94-007-0372-8_9, © Springer Science+Business Media B.V. 2011 451 452 9 A Late Messinian Palynoflora with a Distinct Taphonomy macrofossils collected by Sigurðsson and the palynological content of the sediments. The pollen spectrum presented by Akhmetiev et al. (1978) is short and they reported that over 80% of the pollen was badly deformed and unidentifiable. Pflug (1959) had previously published a pollen list from these sediments but also his sample showed bad preservation and his list is rather incomplete. Based on the literature and our own studies of the macrofossils and sediments in the field it seems likely that both Pflug (1959) and Akhmetiev et al. (1978) obtained their pollen samples from the sandstone/siltstone/lignite fraction of the formation containing wood fragments and leaf impressions. For the present study we analysed pollen from a thin white coloured ash/pumice layer close to the top of the sedimentary succession, positioned a few centimetres above the more oxidized macrofossil layers studied by previous authors. The rich and well-preserved palynoflora from the Selárgil locality contains palaeobiogeographically important elements that can be used to infer routes and modes of plant migration to Iceland during the latest Miocene. 9.2 Geological Setting and Taphonomy The Fnjóskadalur Formation (5.5 Ma; Jancin et al. 1985), North Iceland (Fig. 9.1a, b) is the youngest known Miocene unit yielding plant macrofossils. The formation can be traced along valley sides of the river Fnjóská (Fig. 9.1c), from Mount Reykjafjall to the south, towards the north along the hillsides of Mount Sellandsfjall (including the Selárgil locality; Plate 9.1, 1 and 2), and past the Illugastaðir estate/church site and most of the way down the widening Fnjóskadalur valley past the Ljósavatnsskarð valley crossing (Akhmetiev et al. 1978). Sediments in this region are 25–50 m thick and composed both of clastic and volcanic units (Pjetursson 1905; Sigurðsson 1975; Akhmetiev et al. 1978). The sediments rest on eroded reversely magnetized basalts that are >6.9 Ma (Akhmetiev et al. 1978). The time span represented by the hiatus is uncertain, but basalts covering the sediments (Plate 9.1, 3–5) are between 6 and 5 Ma (Jancin et al. 1985), suggesting a ca 5.5 Ma age for the uppermost fossiliferous part of the underlying sediments. The sedimentary rocks are mostly fluvially originated conglomerates and sandstones, with intercalated lignite and siltstone units in some areas indicating partial lake environments or at least stagnant freshwater. As in other sedimentary sequences in Iceland, a number of volcanic ash layers and tephras/tuff units are found in this formation. The plant macrofossils at the Selárgil locality of the Fnjóskadalur Formation are badly preserved. Most of the sediments containing plant macro-remains (seen below the hammer in Plate 9.1, 5) are red or red-brownish in colour from oxidation, and the fossils are mostly found as impressions (Plate 9.1, 7–10). Faint remnants of coalified material are sometimes visible in the more brownish to greyish samples. The thin white coloured ash/pumice layer topping the macrofossil units (Plate 9.1, 6) and separating them from the fine laminated greyish siltstones just below the overlying basalt, contains no macrofossil. It does, on the other hand, Fig. 9.1 Map showing the fossiliferous locality of the 5.5 Ma formation. (a) Bedrock geology (see Fig. 1.10 for explanation), (b) extension of sedimentary rock formation, (c) Selárgil locality (Geological background modified after Jóhannesson and Sæmundsson 1989; altitudinal lines from Landmælingar Íslands 1990) 454 9 A Late Messinian Palynoflora with a Distinct Taphonomy yield an exceptionally well-preserved palynoflora. Pollen is not abundant in this sample but the preservation is good. Similar excellent preservation was also noticed in the volcanic white ash fall sediments of the 10 Ma sedimentary formation (Tröllatunga locality, see Chap. 6). Interestingly, pollen in the Tertiary formations of Iceland seems to preserve fairly well in very fine ash layers of acid origin (with high silica content) but shows a much worse preservation in the more basic (low silica content) units. It seems likely that pollen contained in this volcanic layer was airborne in the Selárgil region during the actual ash fall and reflects a rather short time interval (few hours, a day to few days) of deposition compared to several of the clastic palynological samples discussed in this book which mostly represent a much longer time interval spanning some years. This might be the reason why several taxa to be expected, such as Juglandaceae, Ulmaceae, etc., were not found in this sample although they occur in both older and younger sediments. Akhmetiev et al. (1978) report some of these taxa (for example, Juglandaceae) in their clastic palynological sample of Selárgil. 9.3 Flora, Vegetation, and Palaeoenvironments The late Messinian flora of Selárgil comprises 53 taxa (Table 9.1, Plates 9.2–9.20) of which by far the most are herbaceous angiosperms (27 taxa; Fig. 9.2). Mosses, ferns and fern allies are represented by seven taxa. Among trees, conifers make up six species and angiosperms ten. Three taxa belong to incertae sedis. Despite the potential taphonomic bias seen in this flora (see above), the vegetation at Selárgil was diverse including wetlands, meadows and well drained lowland and montane forests (Table 9.2, Fig. 9.3). Lowlands were covered by wetlands, rich meadows and shrublands (Fig. 9.4). Stagnant water provided habitats for water plants (Myriophyllum, Nuphar, Menyanthes) and was surrounded by swamp vegetation comprising herbaceous plants and woody shrubs and trees (Ericaceae, Alnus). More closed backswamp forests were probably dominated by Alnus and species of Salix. Well-drained lowland forests including levées and lake margins might have been more diverse in woody species and comprised mixed stands of Betulaceae, Quercus and Salix. Conifers may have been rare elements in the foothill forests but became more abundant in the montane forests where they formed part of mixed broadleaved deciduous and conifer forests (Fig. 9.5). Overall, conifers were quite diverse and may have had different ecologies. For example, Cathaya, which had its last occurrence in Iceland during the deposition of the Fnjóskadalur Formation might have thrived in microclimatically favoured areas, such as humid ravine-like forests, while some others, such as Pinus and Larix, were possibly components of various forest types. Also herbaceous taxa occupied different niches (Table 9.2) as is also seen in the modern vegetation of Iceland. In general, palaeobotanical evidence suggests a typical cool temperate, rather humid, appearance for the late Messinian vegetation in Iceland. A few exotic woody elements persisted from the older floras (Cathaya, Sciadopitys, Tetracentron). 9.3 Flora, Vegetation, and Palaeoenvironments Table 9.1 Taxa recorded for the 5.5 Ma floras of Iceland Fnjóskadalur Formation 5.5 Ma Taxa Bryophyta Sphagnum sp. Equisetaceae Equisetum sp. Lycopodiaceae Lycopodium Polypodiaceae Polypodiaceae gen. et spec. indet. 1 Polypodiaceae gen. et spec. indet. 7 Polypodiaceae gen. et spec. indet. 8 Incertae sedis – unassigned spores Trilete spore, fam., gen. et spec. indet. 2 Pinaceae Abies steenstrupiana Cathaya sp. Picea sect. Picea Pinus sp. 1 (Diploxylon type) Pseudotsuga/Larix sp. Sciadopityaceae Scyadopitys sp. Apiaceae Apiaceae gen. et spec. indet. 6 Apiaceae gen. et spec. indet. 7 Asteraceae Artemisia sp. 2 Asteraceae gen. et spec. indet. 1 Asteraceae gen. et spec. indet. 2 Asteraceae gen. et spec. indet. 4 Betulaceae Alnus cecropiifolia Betula cristata Betula sp. A (section Betulaster) Calycanthaceae aff. Calycanthaceae Caryophyllaceae Caryophyllaceae gen. et spec. indet. 4 Ericaceae Ericaceae gen. et spec. indet. 2 Ericaceae gen. et spec. indet. 3 Fagaceae Quercus infrageneric group Quercus sp. 2 Haloragaceae Myriophyllum sp. 1 Pollen + Leaves RP Other 455 DM 1a + + + + + + + + + + + + + + + + + + + (+) (+) + + + + + + + + + + + 1a 1a 1a 1a 1a 1a 2a 2a 2a 2a 2a 2a 1b 1b 1a 1a 1a 1a 1a, 2a 1a 1a 1b 1b 1b 1b 2b, 3 1b (continued) 456 Table 9.1 (continued) Fnjóskadalur Formation 5.5 Ma Taxa 9 A Late Messinian Palynoflora with a Distinct Taphonomy Pollen Leaves RP Other DM Liliaceae Liliaceae gen. et spec. indet. 4 + 2a Menyanthaceae Menyanthes sp. + 1b Nymphaceae Nuphar sp. + 1b Plantaginaceae aff. Plantago lanceolata + 1b Poaceae Phragmites sp. + 1b Poaceae gen. et spec. indet. 2 + 1b, 2a Poales Poales fam., gen. et spec. indet. + + 1b, 2a Polygonaceae Polygonum viviparum + 1b Ranunculaceae Ranunculus sp. 1 + 1b Ranunculus sp. 2 + 1b Thalictrum sp. 1 + 1b, 2a Ranunculaceae gen. et spec. indet. 2 + 1b Ranunculaceae gen. et spec. indet. 3 + 1b Rosaceae Sanguisorba sp. + 1b, 2a Rosaceae gen. et spec. indet. 10 + 1b Rosaceae gen. et spec. indet. 11 + 1b Rosaceae gen. et spec. indet. 12 + 1b Salicaceae Salix gruberi (+)2 + 1a Salix sp. A (+)2 + 1a Sparganiaceae Sparganium sp. + 1b Trochodendraceae Tetracentron atlanticum + 2a Valerianaceae aff. Valeriana sp. + 1a Incertae sedis – Magnoliophyta Pollen type 21 + ? Pollen type 22 + ? Pollen type 23 + ? L leafy axis, A fruit attached to leafy axis, D fruit dispersed, RP reproductive structure, + organ present, + original description of species based on this organ, (+) organ belonging to genus but uncertain to which of the species, (+) 2 indicating number of pollen types possibly belonging to the eponymous morphotaxon, DM dispersal mode: 1a wind long distance (anemochory), 1b bird long distance (endozoochory), 2a wind short distance (anemochory), 2b animals short distance (exozoochory), 3 dyschory 9.3 Flora, Vegetation, and Palaeoenvironments 457 Fig. 9.2 Distribution of life forms and higher taxa among the plants recovered from the 5.5 Ma sedimentary rock formation. Height of columns indicates number of taxa Fig. 9.3 Schematic block diagram showing palaeo-landscape and vegetation types for the late Late Miocene of Iceland. See Table 9.1 for species composition of vegetation types Table 9.2 Vegetation types and their components during the late Messinian Foothill forests Polypodiaceae gen. et spec. indet. 1, 7, 8 Alnus cecropiifolia Betula cristata Betula sp. A Quercus sp. 2 Betula sp. A Caryophyllaceae gen. et spec. indent. 4 Ericaceae gen. et spec. indet. 2, 3 Plantago lanceolata type Poaceae gen. et spec. indet. 2 Polygonum viviparum Ranunculus sp. 1, 2 Thalictrum sp. 1 Ranunculaceae gen. et spec. indet. 2, 3 Sanguisorba sp. Rosaceae gen. et spec. indet. 10–12 Valerianaceae aff. Valeriana sp. Rocky outcrop forests Lycopodium sp. Pinus sp. 1 Larix sp. Plantago lanceolata type Poaceae gen. et spec. indet. 2 Polygonum viviparum Thalictrum sp. 1 Sanguisorba sp. Rosaceae gen. et spec. indet. 10–12 Tetracentron atlanticum Vegetation types 5.5 Ma Aquatic vegetation Equisetum sp. Myriophyllum sp. 1 Liliaceae gen. et spec. indet. 4 Menyanthes sp. Nuphar sp. Phragmites sp. Sparganium sp. Swamp vegetation Sphagnum sp. Equisetum sp. Apiaceae gen. et spec. indet. 6, 7 Alnus cecropiifolia Ericaceae gen. et spec. indet. 2, 3 Liliaceae gen. et spec. indet. 4 Menyanthes sp. Poaceae gen. et spec. indet. 2 Poales fam. gen. et spec. indet. Sparganium sp. Backswamp forests and temporally flooded lake margin Equisetum sp. Polypodiaceae gen. et spec. indet. 1, 7, 8 Apiaceae gen. et spec. indet. 6, 7 Alnus cecropiifolia aff. Calycanthaceae Poaceae gen. et spec. indet. 2 Salix sp. A Valerianaceae aff. Valeriana sp. Montane forests Abies steenstrupiana Cathaya sp. Picea sp. Levée forests, well-drained lowland forests and lake margins Pinus sp. 1 Polypodiaceae gen. et spec. indet. 1, 7, 8 Larix sp. Alnus cecropiifolia Sciadopitys sp. Betula cristata Betula sp. A aff. Calycanthaceae Tetracentron atlanticum Quercus sp. 2 Salix sp. A Meadows and shrublands Valerianaceae aff. Valeriana sp. Sphagnum sp. Equisetum sp. Lycopodium sp. Apiaceae gen. et spec. indet. 6, 7 Artemisia sp. 2 Asteraceae gen. et spec. indet. 1, 2, 4 AzonAL VeGeTATIon zonAL VeGeTATIon The palaeoecology of fossil species is reconstructed from their sedimentological context and ecology of modern analogues 9.3 Flora, Vegetation, and Palaeoenvironments Fig. 9.4 Schematic transect of a lake margin with moist meadows changing into a light forest dominated by Betulaceae and Salix 459 460 9 A Late Messinian Palynoflora with a Distinct Taphonomy Fig. 9.5 Schematic transect showing of well-drained foothill and montane forest dominated by conifers with admixture of Quercus 9.4 Climatic Requirements of Some Potential Modern Analogues 461 9.4 Climatic Requirements of Some Potential Modern Analogues Cathaya is endemic to central South China (Flora of China Editorial Committee 1999) where it thrives in humid areas between 900 and 1,900 m a. s. l. with MAT 9.3–18.6°C (Cfa climate; Kottek et al. 2006). It typically occurs on slopes and open ridges in connection with mixed mesophytic and broad leaved evergreen forests. Clearly, this genus had a much wider distribution in the past (Liu and Basinger 2000) and persisted in Europe until the Pleistocene. Hence, it may have extended well into cooler variants of humid temperate climate types (Cfb, Cfc climates). Apart from Cathaya, Sciadopitys is the most warmth-loving element among the conifers. At present, Sciadopitys is a monotypic genus (see Chap. 5) confined to cool-temperate, mixed evergreen-deciduous forests, often in pure stands. It thrives in a Cfa to Dfb (snow, fully humid with warm summers; Köppen and Geiger 1928; Kottek et al. 2006) climate with MAT 7.4–16.6°C (temperature range from Utescher & Mosbrugger 2009). Tetracentron (Trochodendraceae) is a monotypic genus with only one living species, Tetracentron sinense Oliv. restricted to central and southwestern China, northern Vietnam, northern Burma, and south of the Himalayas to northeastern India, Bhutan, and eastern Nepal. Tetracentron occurs along streams and forest margins in broadleaved evergreen forests and mixed evergreen-deciduous forests at elevations between 1,100 and 3,500 m a. s. l. (Fu and Bartholomew 2001). It thrives under a variety of climate types (Cfa, Cfb, Cwa, Cwb, Cwc, Dfb; Kottek et al. 2006) with MAT ranging from ca 2.2°C to 19°C. This genus is unambiguously recorded from Icelandic sediments based on diagnostic pollen, fruits, and leaves. It has a stratigraphic range from 15 to 3.6 Ma (see Chap. 12). Another element with a long stratigraphic record is pollen with clear affinities to Calycanthaceae that occur in a similarly wide range of climate types. Quercus is represented by a distinct type of pollen that shows systematic affinities with extant white oaks (infrageneric group Quercus) and red oaks (infrageneric group Lobatae; Denk et al. 2010). Among these groups, the observed vermiculate tectum ornamentation appears to be confined to North American species (Solomon 1983a, b). Among modern oaks, white oaks and red oaks have the most northern and most continental distribution (Camus 1936–1938, 1938–1939, 1952–1954). Red oaks have their centre of diversity in Mexico and Central America but some species can cope with cool temperate climates with winter frosts. The widespread eastern North American Q. rubra L., for example, occurs in humid temperate (Cfa, Cfb, Cfc) and snow (Dfa, Dfb, Dfc) climate types; Kottek et al. 2006) with MAT ranging from −1.1°C to 19.4°C (Thompson et al. 1999). White oaks have a similar range as red oaks in North America but extend even further into cold continental areas with severe winter frosts (Jensen 1997; Nixon and Muller 1997). Quercus macrocarpa Michx. is native to the eastern and mid-western United States and Canada and grows under MAT −1.5°C to 21.8°C (Thompson et al. 1999). 462 The bulk of taxa recorded for the 5.5 Ma formation is not indicative of a particular climate type but rather indifferent and able to thrive in cool and warm temperate climates including snow climates. 9.5 Taxonomic Affinities and origin of newcomers The most spectacular newcomer in the ca 5.5 Ma flora of Iceland is Quercus morphotype 2 with clear affinities to North American white or red oaks (Denk et al. 2010). In North America, red and white oaks extend into areas with MAT below the freezing point and with severe frosts during the winter (see above). Although there is convincing evidence for the formation of glaciers in southern Greenland during the Miocene, these glaciers were confined to mountains at 7.3 Ma (St. John and Krissek 2002), and large-scale northern hemispheric glaciations started not earlier than at the Pliocene-Pleistocene boundary (ca 2.7 Ma, Gibbard and Cohen 2009; East NorwegianGreenland Sea and Barents Sea [Thiede et al. 1998]; 2.8–2.7 Ma Vøring Plateau [Fronval and Jansen 1996]; 2.8–2.6 Ma, Iceland [Geirsdóttir and Eiríksson 1994]; 3.5–2.7 Ma, Greenland [St. John and Krissek 2002]). Plant fossil evidence indicates that Iceland had a warm temperate Cfa climate until at least 12 Ma, and a cool temperate Cfb/Cfc climate suitable for white and/or red oaks until ca 3.6 Ma (see Chap. 13). In view of glaciers of varying size on the mountains of southern Greenland in the Late Miocene and the absence of large-scale ice sheets until the latest Early Pliocene (St. John and Krissek 2002), white and red oaks appear ecologically suited to have colonized Iceland via Greenland from North America during the latest Miocene (ca 6 Ma). Assuming that there is no sampling bias, this type of oak would have migrated to Iceland between 8 and 5.5 Ma (Denk et al. 2010). During the Pliocene and parts of the Early Pleistocene the Earth experienced phases of markedly warm climates. First, between ca 4.5 and 2.7 Ma (Haug et al. 2004), the Mid-Pliocene Climatic Optimum caused conditions in the northern North Atlantic and the Arctic Ocean east of Greenland with summer sea surface temperatures up to >8°C warmer than today (Robinson 2009). In the Early Pleistocene, forest tundra extended to northern Greenland and Bennike (1990) estimated summer temperatures 7–8°C warmer than today. This warming occurred after the first large-scale glaciations in this region (see above) and caused the Inland Ice of Greenland to melt (Bennike 1990). Given such warm conditions in the northern part of Greenland during various times in the later Neogene it appears to be plausible that the link between Greenland and North America via Queen Elizabeth Island may have been passable at the time, when Quercus morphotype 2 migrated to Iceland. This is the last record for the migration of short-distance dispersed plants from the west to Iceland. Among woody plants, two distinct types of Ericaceae tetrads occur for the first time in Iceland in the 5.5 Ma formation (cf. Table 9.2). No closer taxonomic and biogeographic affinities can be established for these types at the moment. The remaining newcomers are widespread cosmopolitan herbaceous taxa with longdistance dispersal and unidentified angiosperms. 9.7 Summary 463 9.6 Comparison to Coeval northern Hemispheric Floras The Late Miocene Lava Camp flora and insect fauna (ca 5.7 ± 0.2 Ma) from the Bering Strait region was described by Hopkins et al. (1971) and Matthews and Ovenden (1990). The flora is dominated by Larix leaves and short shoots. Among the conifers, a member of Pinus subsection Cembrae with a modern Eurasian distribution is noteworthy. The remaining conifer taxa are closely related to species that are at present endemic to northwestern North America (Picea mariana (Mill.) Britton, Sterns & Poggenb., P. glauca (Moench) Voss, P. sitchensis (Bong.) Carrière, Tsuga heterophylla (Raf.) Sarg., T. mertensiana (Bong.) Carrière; Appendix 9.1). In addition, Hopkins et al. (1971) compared cupressaceous pollen to Chamaecyparis that grows as far north as southern Alaska today. Compared to the Icelandic Selárgil flora, the Lava Camp flora was clearly dominated by conifers and, not surprisingly, had closer biogeographic affinities to western North America and Eastern Asia (the Bering land bridge was active until ca 5.5–4.8 Ma; Marincovich and Gladenkov 1999). For the insect assemblage found at Lava Camp, Hopkins et al. (1971) concluded that at present such a fauna could possibly be found in southern British Columbia or northern Washington but not on the modern tundra of Seward Peninsula or in the boreal woodlands of interior Alaska. Today, British Columbia and the northern parts of Washington have humid variants of a Cfb (to Csb) climate, whereas the Bering Strait region (Russian and Alaskan parts) has Dfc (for example, Nome, Alaska, Lieth et al. 1999) or ET climates (Mys Uelen, Russia; Barrow, Alaska; Lieth et al. 1999). In Central Europe, the flora of Murat has been absolutely dated as ca 5.3 Ma (Roiron 1991; Appendix 9.1). This flora is dominated by broadleaved deciduous angiosperms with an admixture of conifers. While most of the genera present in the flora of Murat were also present in the Middle Miocene floras of Iceland (cf. Chaps. 4 and 5), a number of taxa have never been reported from Iceland (Bambusa, Berberis, Cedrela, Celtis, Zelkova). According to Roiron (1991), the lack of Lauraceae, Fagus, Liquidambar, and Platanus, among others, in the flora of Murat along with the greater abundance of temperate and cool elements compared to slightly older floras from the French Massif Central point to a climate cooling in the latest part of the Miocene. 9.7 Summary The Selárgil palynoflora recovered from a thin white coloured ash/pumice layer close to the top of the Fnjóskadalur Formation is ca 5.5 Ma in age. Unlike most other floras from Cainozoic sediments of Iceland, the palynoflora of Selárgil most probably was deposited during a single volcanic eruption. The palynomorphs recorded point to a cool temperate climate (Cfb sensu Köppen) providing suitable conditions for a number of warm-loving relict taxa from older floras of Iceland. Similar warm conditions have been inferred from well-dated more or less coeval sediments in the Bering Strait region that yielded rich plant and insect assemblages. The first appearance in Iceland of a distinct type of Quercus with clear North 464 9 A Late Messinian Palynoflora with a Distinct Taphonomy American biogeographic affinities also indicates that the Greenland-Iceland portion of the North Atlantic Land Bridge was functioning during the Late Miocene. This assumption is based on the fact that acorns of Quercus are not dispersed over long distances by wind or birds. In addition, a Late Miocene migration of Quercus to Iceland from North America would require that suitable (climatic) habitats for oaks extended much further than the Arctic Circle and reached as far north as ca 78°N (Queen Elizabeth Islands). This appears to be plausible in view of various warm phases recorded for the later Cainozoic at high northern latitudes (Mid-Pliocene Climatic Optimum at ca 4.5–2.7 Ma; warm period at ca 2.4–2.1 Ma). During these warm periods, Arctic areas such as northern Greenland experienced conditions with summer sea surface temperatures up to 8°C warmer than today. Hence, colonization of Iceland could have been from northern North America via the Queen Elizabeth Islands, southwards along western Greenland and over a partly emerged Greenland-Iceland ridge. Appendix 9.1 Floristic composition of the 5.5 Ma sedimentary formation of Iceland compared to contemporaneous northern hemispheric fossil assemblages at mid and high-latitudes. Fnjóskadalur flora, Iceland [ca 65°36¢ n, 17°49¢W] 5.5 Ma This study 2 Equisetum sp. 1 Lycopodium 1 Polypodiaceae gen. et spec. indet. 1 1 1 1 1 1, 2 1 1, 3 1 1 1 1, 3 1 1 1 1 1 1 1, 3 Polypodiaceae gen et spec. indet. 7 Polypodiaceae gen. et spec. indet. 8 Sphagnum sp. Trilete spore, fam., gen. et spec. indet. 2 Abies steenstrupiana Cathaya sp. Picea sect. Picea Pinus sp. 1 (Diploxylon type) Pseudotsuga/Larix sp. Scyadopitys sp. Alnus cecropiifolia Apiaceae gen. et spec. indet. 6 Apiaceae gen. et spec. indet. 7 Artemisia sp. 2 Asteraceae gen. et spec. indet. 1 Asteraceae gen. et spec. indet. 2 Asteraceae gen. et spec. indet 4 Betula cristata 3 1 1 1 1 1 1 1 1 2 1 1 3 1 1 1 1 1 1 1 1 1 1 Betula sp. A (section Betulaster) aff. Calycanthaceae Caryophyllaceae gen. et spec. indet. 4 Ericaceae gen. et spec. indet. 2 Ericaceae gen. et spec. indet. 3 Liliaceae gen. et spec. indet. 4 Menyanthes sp. Myriophyllum sp. 1 Nuphar sp. Phragmites sp. aff. Plantago lanceolata Poaceae gen. et spec. indet. 2 Poales fam., gen. et spec. indet. Pollen type 21 Pollen type 22 Pollen type 23 Polygonum viviparum Quercus infrageneric group Quercus sp. 2 Ranunculaceae gen. et spec. indet. 2 Ranunculaceae gen. et spec. indet. 3 Ranunculus sp. 1 Ranunculus sp. 2 Rosaceae gen. et spec. indet. 10 (continued) Appendix 9.1 Fnjóskadalur flora (continued) 1 Rosaceae gen. et spec. indet. 11 1 1, 3 3 1 1 1 1 1 Rosaceae gen. et spec. indet. 12 Salix gruberi Salix sp. A Sanguisorba sp. Sparganium sp. Tetracentron atlanticum Thalictrum sp. 1 aff. Valeriana sp. 3 3 3 3 3 3 3 3 3, 2 3, 2 3 3 Murat flora, France [45°07¢ n, 2°25¢ e] 5.34±0.3 Ma Roiron 1991 2 Abies ramesi 3 Glyptostrobus europaeus 2, 3 Picea sp. 2 Pinus sp. 3 Sequoia langsdorfii 3 3 2 2 3 3 3 3 3 3 3 3 3 2, 3 2, 3 3 3 3 3 3 3 3 3 3 3 Acer campestre Acer integerrimum Acer opulifolium Acer platanoides Acer sanctae-crucis Acer tricuspidatum Alnus glutinosa Alnus hoernesi Alnus sp. cf. A. kefersteinii Alnus viridis Bambusa sp. Berberis sp. cf. B. regeliana Betula sp. Carpinus betulus Carpinus suborientalis Carya minor Cedrela sp. Celtis australis Ceratophyllum demersum cf. Photinia Crataegus sp. cf. C. douglasii Dicotylophyllum sp. 1–5 Dombeyopsis lobata Hedera helix 1 1 1 1, 2 Phellodendron sp. cf. P. amurense Populus tremula Prunus acuminata Quercus hispanica Quercus kubinyi Quercus sp. cf. Q. macranthera Rosa sp. cf. R. californica Tilia tomentosa Ulmus campestris Ulmus sp. cf. U. fulva Zelkova ungeri aff. Z. acuminata Zelkova ungeri aff. Z. crenata 465 Lava Camp flora, Seward Peninsula [65°49¢ n, 163°18¢ W] 5.7 ± 0.2 Ma Hopkins et al. 1971 Matthews and Ovenden 1990 1 Abies 1 Cupressaceae/Taxodiaceae (?Chamaecyparis) 1, 2 Larix sp. (Larix/Pseudotsuga) 1, 2 Picea glauca 1, 2 Picea mariana Picea sitchensis 1, 2 Pinus monticola 1, 2 Pinus subsect. Cembrae 1, 2 Pinus two-needle type undifferentiated Thuja sp. 1 1 1 1 1, 2 1, 2 Tsuga heterophylla Tsuga mertensiana-type Alnus spp. Betula sp. Cornus stolonifera Corylus sp. Carex spp. Cyperus spp. Hippuris sp. Menyanthes trifoliata Onagraceae Paliurus sp. Poaceae Salix sp. Symphoricarpos sp. Ilex sp. aff. I. cornuta 1, 2 Vaccinium sp. Juglans regia 3 Boldface indicates that the genus is present in the 5.5 formation. Grey shading indicates that the genus is present in the younger Tjörnes beds and/or the older Hreðavatn-Stafholt Formation. 1 based on pollen, spores; 2 based on leaves and/or fruit/seed fossils; 3 based on leaf fossils. 466 9 A Late Messinian Palynoflora with a Distinct Taphonomy References Akhmetiev, M. A., Bratzeva, G. M., Giterman, R. E., Golubeva, L. V., & Moiseyeva, A. I. (1978). Late Cenozoic stratigraphy and flora of Iceland. Transactions of the Academy of Sciences USSR, 316, 1–188. Bennike, O. (1990). The Kap København Formation: stratigraphy and palaeobotany of a PlioPleistocene sequence in Peary Land, North Greenland. Meddelelser om Gronland. Geoscience, 23, 1–85. Camus, A. (1936–1938). Les Chênes. Monographie du genre Quercus. Tome I. Genre Quercus, sous-genre Cyclobalanopsis, sous-genre Euquercus (sections Cerris et Mesobalanus). Texte. Paris: Paul Lechevalie. 686 pp. Camus, A. (1938–1939). Les Chênes. Monographie du genre Quercus. Tome II. Genre Quercus, sous-genre Euquercus (sections Lepidobalanus et Macrobalanus). Texte. Paris: Paul Lechevalier. 830 pp. Camus, A. (1952–1954). Les Chênes. Monographie du genre Quercus. Tome III. Genre Quercus: sous-genre Euquercus (sections Protobalanus et Erythrobalanus) et genre Lithocarpus. Texte. Paris: Paul Lechevalier. 1314 pp. Denk, T., Grímsson, F., & Zetter, R. (2010). Episodic migration of oaks to Iceland: Evidence for a North Atlantic “land bridge” in the latest Miocene. American Journal of Botany, 97, 276–287. Flora of China Editorial Committee. (1999). Flora of China, Cycadaceae through Fagacaeae (Vol. 4). St. Louis: Missouri Botanical Garden Press. 453 pp. Fronval, T., & Jansen, E. (1996). Late Neogene paleoclimates and paleoceanography in the Iceland-Norwegian Sea: Evidence from the Iceland and Vøring Plateaus. Proceedings of the Ocean Drilling Program. Scientific Results 151, 455–468. Fu, D., & Bartholomew, B. (2001). Tetracentraceae. In Editorial Committee of the Flora of China (Ed.), Flora of China, Caryophyllaceae through Lardizabalaceae (Vol. 6, p. 125). St. Louis: Missouri Botanical Garden Press. Geirsdóttir, Á., & Eiríksson, J. (1994). Growth of an intermittent ice sheet in Iceland during the Late Pliocene and Early Pleistocene. Quaternary Research, 42, 115–130. Gibbard, P. L., & Cohen, K. M. (2009). Global chronostratigraphical correlation table for the last 2.7 million years. v. 2009. http://www.quaternary.stratigraphy.org.uk/charts/. Haug, G. H., Tiedemann, R., & Keigwin, L. D. (2004). How the Isthmus of Panama put ice in the Arctic. Oceanus, 42(2), 1–4. Hopkins, D. M., Matthews, J. V., Wolfe, J. A., & Silberman, M. L. (1971). A Pliocene flora and insect fauna from the Bering Strait region. Palaeogeography, Palaeoclimatology, Palaeoecology, 9, 211–231. Jancin, M., Young, K., Voight, B., Aronson, J., & Saemundsson, K. (1985). Stratigraphy and K/ Ar ages across the west flank of the Northeast Iceland axial rift zone, in relation to the 7 Ma volcano-tectonic reorganization of Iceland. Journal of Geophysical Research, 90(B12), 9961–9985. Jensen, R. J. (1997). Quercus Linnaeus sect. Lobatae Loudon, Hort. Brit., 385. 1830. Red or black oaks. In Flora of North America Editorial Committee (Ed.), Flora of North America North of Mexico, Magnoliophyta: Magnoliidae and Hamamelidae (Vol. 3, pp. 447–468). New York: Oxford University Press. Jóhannesson, H., & Sæmundsson, K. (1989). Geological map of Iceland. 1:500, 000. Bedrock Geology (1st ed.). Reykjavík: Icelandic Museum of Natural History/Icelandic Geodetic Survey. St John, K. E. K., & Krissek, L. A. (2002). The late Miocene to Pleistocene ice-rafting history of southeast Greenland. Boreas, 31, 28–35. Köppen, W., & Geiger, R. (1928). Klimakarte der Erde, Wall-map 150 cm × 200 cm. Gotha: Verlag Justus Perthes. Kottek, M., Grieser, J., Beck, C., Rudolf, B., & Rubel, F. (2006). World map of the Köppen-Geiger climate classification updated. Meteorologische Zeitschrift, 15, 259–263. Explanation of Plates 467 Landmælingar Íslands. (1990). Uppdráttur Íslands. Blað 73, Lundabrekka. Scale 1:100000. Liu, Y.-S., & Basinger, J. F. (2000). Fossil Cathaya from the Canadian High Arctic. International Journal of Plant Sciences, 161, 829–847. Marincovich, L., Jr., & Gladenkov, A. Y. (1999). Evidence for an early opening of the Bering Strait. Nature, 397, 149–151. Matthews, J. F., Jr., & Ovenden, L. E. (1990). Late tertiary plant macrofossils from localities in Arctic/Subarctic North America: A review of the data. Arctic, 43, 364–392. Nixon, K. C., & Muller, C. H. (1997). Quercus Linnaeus sect. Quercus. White oaks. In Flora of North America Editorial Committee (Ed.), Flora of North America North of Mexico, Magnoliophyta: Magnoliidae and Hamamelidae, (Vol. 3). New York: Oxford University Press. 471–506 pp. Pflug, H. D. (1959). Sporenbilder aus Island und ihre stratigraphische Deutung. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen, 107, 141–172. Pjetursson, H. (1905). Om Islands Geologi. Meddelelser fra Dansk Geologisk Førening, 2(11), 1–104. Robinson, M. M. (2009). New quantitative evidence of extreme warmth in the Pliocene Arctic. Stratigraphy, 6, 265–275. Roiron, P. (1991). La macroflore d’age Miocene superieur des diatomites de Murat (Cantal, France) Implications paleoclimatiques. Palaeontographica B, 223, 169–203. Sigurðsson, O. (1975). Steingervingar í Selárgili í Fnjóskadal. Týli, 5, 1–6. Solomon, A. M. (1983a). Pollen morphology and plant taxonomy of white oaks in eastern North America. American Journal of Botany, 70, 481–494. Solomon, A. M. (1983b). Pollen morphology and plant taxonomy of red oaks in eastern North America. American Journal of Botany, 70, 495–507. Thiede, J., Winkler, A., Wolfwelling, T., Eldholm, O., Myhre, A. M., Baumann, K. H., Henrich, R., & Stein, R. (1998). Late Cenozoic history of the polar North Atlantic – results from ocean drilling. Quaternary Science Reviews, 17, 185–208. Thompson, R. S., Anderson, K. H., & Bartlein, P. J. (1999). Atlas of relations between climatic parameters and distribution of important trees and shrubs in North America-Hardwoods. U.S. Geological Survey Professional Paper, 1650-B, 1–423. Utescher, T., & Mosbrugger, V. (2009). Palaeoflora Database. http://www.geologie.unibonn.de/ Palaeoflora explanation of Plates Plate 9.1 1. Selárgil in Fnjóskadalur, Fnjóskadalur Formation (ca 5.5 Ma). View up the gully Selárgil, outcrop seen in the distance to the left. 2. View over the upper part of the Selárgil gully, outcrop below the massive columnar basalt. 3. Selárgil outcrop, geologist digging for fossils. 4. Upper part of the sedimentary section, showing the brown sandy siltstones. 5. Contact zone between basalt and sediments. 6. Detail showing fine grained clay-rich siltstones and the white tephra layer at the bottom were the pollen originate from. 7–10. Preservation of fossils, compressions and impressions in siltstone (7, 9), and fine grained sandstone (8), and a lignified part of stem. Reddish colour caused due to oxidization by weathering Plate 9.2 1–3. Sphagnum sp. 1. Spore in SEM, distal polar view. 2. Detail of spore surface. 3. Spore in LM, proximal polar view showing trilete tetrad mark. 4–6. Sphagnum sp. 4. Spore in SEM, proximal polar view. 5. Detail of spore surface. 6. Spore in LM, proximal polar view 468 9 A Late Messinian Palynoflora with a Distinct Taphonomy showing trilete tetrad mark. 7–9. Lycopodium sp. 7. Spore in SEM, distal polar view. 8. Detail of spore surface. 9. Spore in LM, oblique polar view. 10–12. Lycopodium sp. 10. Spore in SEM, proximal polar view showing trilete tetrad mark. 11. Detail of spore surface. 12. Spore in LM, polar view Plate 9.3 1–3. Polypodiaceae gen. et spec. indet. 1. 1. Spore in SEM, equatorial view showing monolete tetrad mark. 2. Detail of spore surface. 3. Spore in LM, equatorial view. 4–6. Polypodiaceae gen. et spec. indet. 7. 4. Spore in SEM, equatorial view. 5. Detail of spore surface. 6. Spore in LM, equatorial view. 7–9. Polypodiaceae gen. et spec. indet. 8. 7. Spore in SEM, oblique equatorial view. 8. Detail of spore surface. 9. Spore in LM, oblique equatorial view. 10–12. Trilete spore fam. gen. et spec. indet. 2. 10. Spore in SEM, distal polar view. 11. Detail of spore surface. 12. Spore in LM, proximal polar view showing trilete tetrad mark Plate 9.4 1. Equisetum sp., underground rhizome (IMNH org 65-02). 2. Equisetum sp., detail showing part of aerial stem (IMNH org 67). 3. Equisetum sp., part of stem (IMNH 4326-03). 4. Equisetum sp., detail showing sheath (IMNH org 67). 5. Abies steenstrupiana, cone scale (IMNH org 65-01). 6. Picea sect. Picea, needle (IMNH org 62-02). 7. Poales fam. gen. et spec. indet. 2. (IMNH 6823). 8. Phragmites sp., part of stem (IMNH 4325-02) Plate 9.5 1–3. Abies sp. 1. Bisaccate pollen grain in SEM, proximal polar view. 2. Detail of corpus surface. 3. Bisaccate pollen grain in LM, polar view. 4. Pinus sp. 1 (Diploxylon type), bisaccate pollen grain, equatorial view. 5. Cathaya sp., bisaccate pollen grain, polar view. 6–8. Larix/Pseudotsuga sp. 6. Pollen grain in SEM. 7. Detail of pollen grain surface. 8. Pollen grain in LM. 9–11. Sciadopitys sp. 9. Pollen grain in SEM, distal polar view. 10. Detail of pollen grain surface. 11. Pollen grain in LM Plate 9.6 1–3. Apiaceae gen. et spec. indet. 6. 1. Pollen grain in SEM, equatorial view. 2. Detail of pollen grain surface. 3. Pollen grain in LM, equatorial view. 4–6. Apiaceae gen. et spec. indet. 7. 4. Pollen grain in SEM, equatorial view. 5. Detail of pollen grain surface. 6. Pollen grain in LM, equatorial view. 7–10. Artemisia sp. 2. 7. Pollen grain in SEM, equatorial view. 8. Detail of pollen grain surface. 9. Pollen grain in LM, polar view (upper), equatorial view (lower). 10–12. Artemisia sp. 2. 10. Pollen grain in SEM, oblique view. 11. Detail of pollen grain surface. 12. Pollen grain in LM, polar view Plate 9.7 1–3. Asteraceae gen. et spec. indet. 1 (Liguliflorae) 1. Pollen grain in SEM. 2. Detail of pollen grain surface. 3. Pollen grain in LM. 4–6. Asteraceae gen. et spec. indet. 2. 4. Pollen grain in SEM, polar view. 5. Detail of pollen grain surface. 6. Pollen grain in LM, polar view. 7–9. Asteraceae gen. et spec. indet 4. 7. Pollen grain in SEM. 8. Detail of pollen grain surface. 9. Pollen grain in LM Plate 9.8 1. Alnus cecropiifolia, large wide ovate leaf (IMNH 4333) 2. Betula cristata, lower part of leaf, cordate base (IMNH 4332) Plate 9.9 1. Betula sp. A (section Betulaster) (IMNH 4339-02). 2. Detail of Fig. 1 showing teeth along margin Plate 9.10 1–3. Alnus sp. 1. 1. Pollen grain in SEM, polar view. 2. Detail of pollen grain surface. 3. Pollen grain in LM, polar view. 4–6. Betula sp. 4. Pollen grain in SEM oblique equatorial view. 5. Detail of pollen grain surface. 6. Pollen grain in LM, polar view. 7–9. Betula sp. 7. Pollen grain in SEM, polar view. 8. Detail of pollen grain surface. 9. Pollen grain in LM, polar view. 10. Betula sp., pollen grain in LM, polar view. 11. Betula sp., pollen grain in LM, polar view Explanation of Plates 469 Plate 9.11 1–3. aff. Calycanthaceae sp. 1. Pollen grain in SEM, equatorial view. 2. Detail of pollen grain surface. 3. Pollen grain in LM, equatorial view. 4–6. Caryophyllaceae gen. et spec. indet. 4. 4. Pollen grain in SEM. 5. Detail of pollen grain surface. 6. Pollen grain in LM. 7–9. Ericaceae gen. et spec. indet. 2. 7. Tetrad in SEM. 8. Detail of tetrad surface. 9. Tetrad in LM Plate 9.12 1–3. Ericaceae gen. et spec. indet. 3. 1. Tetrad in SEM. 2. Detail of tetrad surface. 3. Tetrad in LM. 4–6. Quercus infrageneric group Quercus sp. 2. 4. Pollen grain in SEM, equatorial view. 5. Detail of pollen grain surface. 6. Pollen grain in LM, equatorial view. 7–9. Quercus infrageneric group Quercus sp. 2. 7. Pollen grain in SEM, equatorial view. 8. Detail of pollen grain surface. 9. Pollen grain in LM, equatorial view. 10–12. Myriophyllum sp. 1. 10. Pollen grain in SEM, oblique polar view. 11. Detail of pollen grain surface. 12. Pollen grain in LM, polar view Plate 9.13 1–3. Liliaceae gen. et spec. indet. 4. 1. Pollen grain in SEM. 2. Detail of pollen grain surface. 3. Pollen grain in LM. 4–6. Menyanthes sp. 4. Pollen grain in SEM, equatorial view. 5. Detail of pollen grain surface. 6. Pollen grain in LM, equatorial view. 7–9. Nuphar sp. 7. Pollen grain in SEM. 8. Detail of pollen grain surface. 9. Pollen grain in LM. 10–12. Plantago lanceolata. 10. Pollen grain in SEM. 11. Detail of pollen grain surface. 12. Pollen grain in LM Plate 9.14 1–3. Poaceae gen. et spec. indet. 2. 1. Pollen in SEM. 2. Detail of pollen grain surface. 3. Pollen grain in LM. 4–6. Polygonum viviparum. 4. Pollen grain SEM, equatorial view. 5. Detail of pollen grain surface, polar area. 6. Pollen grain in LM, equatorial view. 7–9. Ranunculus sp. 1. 7. Pollen grain in SEM. 8. Detail of pollen grain surface. 9. Pollen grain in LM. 10–12. Ranunculus sp. 2. 10. Pollen grain in SEM, equatorial view. 11. Detail of pollen grain surface. 12. Pollen grain in LM, equatorial view Plate 9.15 1–3. Thalictrum sp. 1. 1. Pollen grain in SEM. 2. Detail of pollen grain surface. 3. Pollen grain in LM. 4–6. Ranunculaceae gen. et spec. indet 2. 4. Pollen grain in SEM, equatorial view. 5. Detail of pollen grain surface. 6. Pollen grain in LM, equatorial view. 7–9. Ranunculaceae gen. et spec. indet 2. 7. Pollen grain in SEM, equatorial view. 8. Detail of pollen grain surface. 9. Pollen in LM, equatorial view. 10–12. Ranunculaceae gen et spec. indet 3. 10. Pollen grain in SEM, equatorial view. 11. Detail of pollen grain surface. 12. Pollen grain in LM, equatorial view Plate 9.16 1–3. Sanguisorba sp. 1. Pollen grain in SEM, polar view. 2. Detail of pollen grain surface. 3. Pollen grain in LM, polar view. 4–6. Sanguisorba sp. 4. Pollen grain in SEM, equatorial view. 5. Detail of pollen grain surface. 6. Pollen grain in LM, equatorial view. 7–9. Rosaceae gen. et spec. indet. 10. 7. Pollen in SEM, equatorial view. 8. Detail of pollen grain surface. 9. Pollen grain in LM, equatorial view. 10–12. Rosaceae gen. et spec. indet. 10. 10. Pollen grain in SEM, equatorial view. 11. Detail of pollen grain surface. 12. Pollen grain in LM, equatorial view Plate 9.17 1–3. Rosaceae gen. et spec. indet. 11. 1. Pollen grain in SEM, equatorial view. 2. Detail of pollen grain surface. 3. Pollen grain in LM, equatorial view. 4–6. Rosaceae gen. et spec. indet. 12. 4. Pollen grain in SEM, equatorial view. 5. Detail of pollen grain surface. 6. Pollen grain in LM, equatorial view. 7–9. Salix sp. 4. 7. Pollen grain in SEM, equatorial view. 8. Detail of pollen grain surface. 9. Pollen grain in LM, equatorial view. 10–12. Salix sp. 5. 10. Pollen grain in SEM, equatorial view. 11. Detail of pollen grain surface. 12. Pollen grain in LM, equatorial view Plate 9.18 1. Salix gruberi, lower part of large leaf (IMNH org 63-01). 2. Detail of Fig. 1 showing venation and teeth along margin. 3. Salix sp. A, narrow elliptic leaf (IMNH 4326-01). 4. Detail of Fig. 3 showing venation along margin. 5. Salix gruberi, upper half of leaf (IMNH 4325-01). 6. Salix gruberi, lower part of leaf (IMNH 4324) 470 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.19 1–3. Sparganium sp. 1. Pollen grain in SEM. 2. Detail of pollen grain surface. 3. Pollen grain in LM. 4–6. Tetracentron atlanticum. 4. Pollen grain in SEM, equatorial view. 5. Detail of pollen grain surface. 7–9. aff. Valeriana sp. 7. Pollen grain in SEM, equatorial view. 8. Detail of pollen grain surface. 9. Pollen in LM, polar view Plate 9.20 1–3. Pollen type 21. 1. Pollen grain in SEM, equatorial view. 2. Detail of pollen grain surface. 3. Pollen grain in LM, polar view (left), equatorial view (right). 4–6. Pollen type 22. 4. Pollen grain in SEM, equatorial view. 5. Detail of pollen grain surface. 6. Pollen grain in LM, equatorial view. 7–9. Pollen type 23. 7. Pollen grain in SEM, equatorial view. 8. Detail of pollen grain surface. 9. Pollen grain in LM, equatorial view Plates Plate 9.1 472 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.2 Plates 473 Plate 9.3 474 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.4 Plates 475 Plate 9.5 476 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.6 Plates 477 Plate 9.7 478 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.8 Plates 479 Plate 9.9 480 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.10 Plates 481 Plate 9.11 482 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.12 Plates 483 Plate 9.13 484 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.14 Plates 485 Plate 9.15 486 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.16 Plates 487 Plate 9.17 488 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.18 Plates 489 Plate 9.19 490 9 A Late Messinian Palynoflora with a Distinct Taphonomy Plate 9.20