(Table 2) Survivorship of transplanted seedlings of Picea glauca inside and outside three tree islands in 1994, 1995, 2009 and after 15 years

The northern boundary of boreal forest and the ranges of tree species are expected to shift northward in response to climate warming, which will result in a decrease in the albedo of areas currently covered by tundra vegetation, an increase in terrestrial carbon sequestration, and an alteration of biodiversity in the current Low Arctic. Central to the prediction of forest expansion is an increase in the reproductive capacity and establishment of individual trees. We assessed cone production, seed viability, and transplanted seedling success of Picea glauca (Moench.) Voss. (white spruce) in the early 1990s and again in the late 2000s at four forest stand sites and eight tree island sites (clonal populations beyond present treeline) in the Mackenzie Delta region of the Northwest Territories, Canada. Over the past 20 years, average temperatures in this region have increased by 0.9 °C. This area has the northernmost forest-tundra ecotone in North America and is one of the few circumpolar regions where the northern limit of conifer trees reaches the Arctic Ocean. We found that cone production and seed viability did not change between the two periods of examination and that both variables decreased northward across the forest-tundra ecotone. Nevertheless, white spruce individuals at the northern limit of the forest-tundra ecotone produced viable seeds. Furthermore, transplanted seedlings were able to survive in the northernmost sites for 15 years, but there were no signs of natural regeneration. These results indicate that if climatic conditions continue to ameliorate, reproductive output will likely increase, but seedling establishment and forest expansion within the forest-tundra of this region is unlikely to occur without the availability of suitable recruitment sites. Processes that affect the availability of recruitment sites are likely to be important elsewhere in the circumpolar ecotone, and should be incorporated into models and predictions of climate change and its effects on the northern forest-tundra ecotone.

Palynology and radiocarbon dating on sediment profile PG1756 from Lake Billyakh, NE Siberia, Russia

In this study a radiocarbon-dated pollen record from Lake Billyakh (65°17'N, 126°47'E; 340 m a.s.l.) in the Verkhoyansk Mountains was used to reconstruct vegetation and climate change since about 15 kyr BP (1 kyr=1000 cal. yr). The pollen record and pollen-based biome reconstruction suggest that open cool steppe and grass and sedge tundra communities with Poaceae, Cyperaceae, Artemisia, Chenopodiaceae, Caryophyllaceae and Selaginella rupestris dominated the area from 15 to 13.5 kyr BP. On the other hand, the constant presence of Larix pollen in quantities comparable to today's values points to the constant presence of boreal deciduous conifer trees in the regional vegetation during the last glaciation. A major spread of shrub tundra communities, including birch (Betula sect. Nanae), alder (Duschekia fruticosa) and willow (Salix) species, is dated to 13.5-12.7 kyr BP, indicating a noticeable increase in precipitation toward the end of the last glaciation, particularly during the Allerød Interstadial. Between 12.7 and 11.4 kyr BP pollen percentages of herbaceous taxa rapidly increased, whereas shrub taxa percentages decreased, suggesting strengthening of the steppe communities associated with the relatively cold and dry Younger Dryas Stadial. However, the pollen data in hand indicate that Younger Dryas climate was less severe than the climate during the earlier interval from 15 to 13.5 kyr BP. The onset of the Holocene is marked in the pollen record by the highest values of shrub and lowest values of herbaceous taxa, suggesting a return of warmer and wetter conditions after 11.4 kyr BP. Percentages of tree taxa increase gradually and reach maximum values after 7 kyr BP, reflecting the spread of boreal cold deciduous and taiga forests in the region. An interval between 7 and 2 kyr BP is noticeable for the highest percentages of Scots spine (Pinus subgen. Diploxylon), spruce (Picea) and fir (Abies) pollen, indicating mid-Holocene spread of boreal forest communities in response to climate amelioration and degradation of the permafrost layer.

A new modern analogue reference dataset and its application to the 430-kyr pollen record from Lake Biwa

This study presents a newly compiled dataset of modern pollen and climate data from 798 sites across Japan and the Russian Far East. This comprehensive reference dataset combined with the modern analogue technique (MAT) provides a powerful tool for pollen-based reconstruction of the Quaternary Northwest Pacific climate. Pollen-derived reconstruction of the modern climate at the reference pollen-sampling sites matches well with the estimated modern climate values (R2 values vary between 0.79 and 0.95, and RMSEP values vary between 5.8 and 9.7% of the modern climatic range for all nine tested variables). The successful testing of the method encourages its application to the fossil pollen records. We used a coarse-resolution pollen record from Lake Biwa to reconstruct glacial-interglacial climate dynamics in central Japan since ~438 kyr and compared it to the earlier reconstruction based on a less representative reference dataset. The current and earlier results consistently demonstrate that the coldest glacial intervals experienced pronounced cooling in winter and moderate cooling in summer, supporting the growth of cool mixed forest (COMX) where warm mixed forest (WAMX) predominates today. During the last glacial, maximum (~24 kyr BP) mean temperatures of the coldest (MTCO) and warmest (MTWA) month were about -13 °C (RMSEP = 2.34 °C) and 21 °C (RMSEP = 1.66 °C) respectively, and annual precipitation (PANN) was about 800 mm (RMSEP = 158.06 mm). During the thermal optimums of the interglacial intervals, the temperatures of the coldest and warmest month were above 0 °C and 25 °C respectively, leading to the reconstruction of WAMX and temperate conifer forest (TECO). Although both these vegetation types grow in the southern part of Japan today, WAMX requires warmer space. The presence of WAMX during marine isotope stages (MIS) 11 and 1, and its absence during MIS 9 and MIS 5 contradict the marine isotope and Antarctic ice records, suggesting that the latter two interglacials were the warmest of the last 800 kyr. The apparent contradiction allows at least three different explanations including low temporal resolution of the pollen record; different trends in CO2 concentrations during 'short' and 'long' interglacials; and regional climate variability and non-linear response of different regions to the global forcing. More definitive conclusions will be possible on the basis of forthcoming high-resolution pollen records from central Japan.

Sporomorphs and dinoflagellate cysts of ODP Leg 113 holes

Palynological studies were carried out on Paleogene sections from Sites 693 and 696 of Ocean Drilling Project Leg 113 in the Weddell Sea region. Dinoflagellate cysts and sporomorphs were recovered at Site 696 (61°S, 42°W) indicating a middle Eocene to late Eocene/earliest Oligocene age for a glauconitic silt/sandstone. At Site 693 (70°S, 14°W) early Oligocene siliciclastic mud contains a low diversity palynoflora. In an upper Oligocene section (Site 693) only rare, reworked Mesozoic palynomorphs were encountered. Palynological data from Kerogen analyses, dinocysts, and sporomorphs are used to reconstruct the climatic change on the South Orkney microcontinent from the middle Eocene to the late Eocene/earliest Oligocene at Site 696 and the late early Oligocene/early late Oligocene time interval at Site 693 near the continental margin. The middle Eocene was a warm period in the Orkney region with good growing conditions for a warm temperate Nothofagus/conifer forest with an admixture of Proteaceae. Temperate surface water masses, which allowed the growth of a reasonably diverse dinocyst assemblage (ca. 15-20 species), persisted until the end of the Eocene at Site 696. Late early Oligocene sediments of Site 693 (Antarctic continental margin) contain only a low diversity dinocyst flora (two species). The major Cenozoic cooling event in the Weddell Sea region probably occurred at the Eocene/Oligocene boundary. A second dramatic climatic deterioration seems to have taken place during the late early/early late Oligocene, when dinocysts disappeared at the Dronning Maud Land margin area.

Dinoflagellates and pollen of the Brito-Arctic Igneous Province flora, ODP Hole 104-642E

Palynological studies of the intrabasaltic sediment layers in the lower volcanic series from ODP Leg 104 outer Voring Plateau Hole 642E Cores 102 through 109 indicated abundant pollen and rarer dinoflagellate cysts. The dinoflagellates belong to the Apectodinium hyperacanthum Zone and indicate an age equivalent to nannoplankton Zones NP9-lower NP10 around the Paleocene/Eocene boundary. The pollen and spore assemblage found here in 12 of the samples from the lower volcanic series is of well- preserved and distinctive specimens and contains unusual forms of pollen from the Taxodiaceae and the Hamamelidae. It has not been transported far from vegetation that was dominated by conifer forest with some ferns and deciduous arborescent angiosperms. Nearly identical assemblages are found elsewhere in the Brito-Arctic Igneous Province, in intrabasaltic sediments from eastern Greenland, the Faeroe Islands, the Isle of Mull, and Antrim (Northern Ireland), and above basalt at the Rockall Plateau. The assemblage is also present in sediments around the Paleocene/Eocene boundary in Spitsbergen. This pollen and spore flora is also associated with dinoflagellate cysts of the Apectodinium hyperacanthum Zone in the deposits from eastern Greenland, the Rockall Plateau, and Spitsbergen, suggesting that these are correlative. Assemblages of the same age from the North Sea, Denmark, and the London and Paris Basins are different. Paleobotanical evidence suggests a short survival of the intrabasaltic flora, and that all the deposits considered here are of about the same age. We propose that at around the Paleocene/Eocene boundary a distinct flora, named here as the Brito-Arctic Igneous Province (BIP) flora, occurred on the line of volcanicity stretching from Rockall to the Greenland Sea, and even to Spitsbergen. Geophysical evidence supports our view that the Rockall to East Greenland intrabasaltics are more or less contemporaneous, at about the Paleocene/Eocene boundary. However, the comparable pollen and spore assemblage in the Hebridean province, at Mull and Antrim, is from pyroclastics that may be a little older.

Palynology, source vegetation, environment and age of ODP Site 188-1166 sediments

Twenty-three core catcher samples from Site 1166 (Hole 1166A) in Prydz Bay were analyzed for their palynomorph content, with the aims of determining the ages of the sequence penetrated, providing information on the vegetation of the Antarctic continent at this time, and determining the environments under which deposition occurred. Dinocysts, pollen and spores, and foraminiferal test linings were recovered from most samples in the interval from 142.5 to 362.03 meters below seafloor (mbsf). The interval from 142.5 to 258.72 mbsf yielded palynomorphs indicative of a middle-late Eocene age, equivalent to the lower-middle Nothofagidites asperus Zone of the Gippsland Basin of southeastern Australia. The Prydz Bay sequence represents the first well-dated section of this age from East Antarctica. Dinocysts belonging to the widespread "Transantarctic Flora" give a more confident late Eocene age for the interval 142.5-220.5 mbsf. The uppermost two cores within this interval, namely, those from 142.5 and 148.36 mbsf, show significantly higher frequencies of dinocysts than the cores below and suggest that an open marine environment prevailed at the time of deposition. The spore and pollen component may reflect a vegetation akin to the modern rainforest scrubs of Tasmania and New Zealand. Below 267 mbsf, sparse microfloras, mainly of spores and pollen, are equated with the Phyllocladidites mawsonii Zone of southeastern Australia, which is of Turonian to possibly Santonian age. Fluvial to marginal marine environments of deposition are suggested. The parent vegetation from this interval is here described as "Austral Conifer Woodland." The same Late Cretaceous microflora occurs in two of the cores above the postulated unconformity at 267 mbsf. In the core at 249.42 mbsf, the Late Cretaceous spores and pollen are uncontaminated by any Tertiary forms, suggesting that a clast of this older material has been sampled; such a clast may reflect transport by ice during the Eocene. At 258.72 mbsf, Late Cretaceous spores and pollen appear to have been recycled into the Eocene sediments.

(Table 1) Comparison of localities where Switzianthus (Anderson & Anderson) has been found with Dejerseya and Heidiphyllum occurrence

Fossil leaves of the Voltziales, an ancestral group of conifers, rank among the most common plant fossils in the Triassic of Gondwana. Even though the foliage taxon Heidiphyllum has been known for more than 150 years, our knowledge of the reproductive organs of these conifers still remains very incomplete. Seed cones assigned to Telemachus have become increasingly well understood in recent decades, but the pollen cones belonging to these Mesozoic conifers are rare. In this contribution we describe the first compression material of a voltzialean pollen cone from Upper Triassic strata of the Transantarctic Mountains. The cone can be assigned to Switzianthus Anderson & Anderson, a genus that was previously assumed to belong to an enigmatic group of pteridosperms from the Triassic Molteno Formation of South Africa. The similarities of cuticle and pollen morphology, together with co-occurrence at all known localities, indicate that Switzianthus most probably represents the pollen organ of the ubiquitous Heidiphyllum/Telemachus plant.

Early Albian palynoflora of ODP Hole 120-750B

A well-preserved, diverse sporomorph flora of over 60 species has been found in Cores 120-750B-12W through -14R from the Southern Kerguelen Plateau. Analysis of the flora indicates that the terrestrial sediments overlaying the basaltic basement are late Early Cretaceous in age. Ranges of the sporomorphs in other parts of Gondwana and the morphology and paucity of angiosperm pollen grains confine the age of this section to the early to possibly early middle Albian. The Albian palynomorph assemblages in Hole 750B are composed primarily of fern spores and podocarpaceous pollen, and show most similarity to those from southern Australia. Changes in the flora through time reflect the successional vegetation changes on barren volcanic islands, beginning with high percentages of colonizing ferns and maturing into conifer (podocarp) forests. The flora shows some signs of endemism, which may be a result of the isolated position of the Kerguelen Islands during the Early Cretaceous. This endemism is expressed by high percentages of a distinctive monosulcate pollen species Ashmoripollis woodywisei n.sp. of pteridosperm or cycadophytean origin, and by a thick-walled, monosulcate angiosperm pollen species of the genus Clavatipollenites. The climatic conditions were probably cool to temperate (mean annual temperature approximately 7°-12°C) and humid (annual rainfall >1000 mm), analogous to modern Podocarpus-dominated forests in New Zealand and in South American mountain regions.

Results of n-alkane biomarker analyses and paleoclimate reconstruction

The current study presents quantitative reconstructions of tree cover, annual precipitation and mean July temperature derived from the pollen record from Lake Billyakh (65°17'N, 126°47'E, 340 m above sea level) spanning the last ca. 50 kyr. The reconstruction of tree cover suggests presence of woody plants through the entire analyzed time interval, although trees played only a minor role in the vegetation around Lake Billyakh prior to 14 kyr BP (<5%). This result corroborates low percentages of tree pollen and low scores of the cold deciduous forest biome in the PG1755 record from Lake Billyakh. The reconstructed values of the mean temperature of the warmest month ~8-10 °C do not support larch forest or woodland around Lake Billyakh during the coldest phase of the last glacial between ~32 and ~15 kyr BP. However, modern cases from northern Siberia, ca. 750 km north of Lake Billyakh, demonstrate that individual larch plants can grow within shrub and grass tundra landscape in very low mean July temperatures of about 8 °C. This makes plausible our hypothesis that the western and southern foreland of the Verkhoyansk Mountains could provide enough moist and warm microhabitats and allow individual larch specimens to survive climatic extremes of the last glacial. Reconstructed mean values of precipitation are about 270 mm/yr during the last glacial interval. This value is almost 100 mm higher than modern averages reported for the extreme-continental north-eastern Siberia east of Lake Billyakh, where larch-dominated cold deciduous forest grows at present. This suggests that last glacial environments around Lake Billyakh were never too dry for larch to grow and that the summer warmth was the main factor, which limited tree growth during the last glacial interval. The n-alkane analysis of the Siberian plants presented in this study demonstrates rather complex alkane distribution patterns, which challenge the interpretation of the fossil records. In particular, extremely low n-alkane concentrations in the leaves of local coniferous trees and shrubs suggest that their contribution to the litter and therefore to the fossil lake sediments might be not high enough for tracing the Quaternary history of the needleleaved taxa using the n-alkane biomarker method.

Palynology of ODP Leg 127 sites

This initial survey of pollen from 192 samples from Hole 794A, supplemented by 189 samples from Hole 795 and 797B, suggests that marine pollen assemblages from the southwestern Sea of Japan provide a consistent Neogene pollen stratigraphy and a solid basis for regional paleoenvironmental reconstructions. Late Miocene vegetation inferred from these pollen data, a mix of conifer and broad-leaf elements with now-extinct Tertiary types well represented, appears similar to Aniai-type floras of Japan. During the late Miocene through early Pliocene, as Tertiary types declined, conifers (including the Sequoia/Cryptomeria group) became more prominent than broad-leaf elements, and herbs played an increasing role in the vegetation. Middle Pliocene pollen assemblages imply significant changes in forest composition. In a 500,000-yr interval centered at ~4 m.y., Tertiary and warm-temperate deciduous types re-expanded and were comparable to or greater than middle-late Miocene levels. Temperate and cold-temperate conifers {Picea, Abies, Tsuga) were minimal. Subsequently, Tertiary and deciduous forest components (including Quercus) decreased, Picea, Tsuga, and Abies were again prominent, and herbs formed an increasingly larger part of the vegetation. Between ~3 m.y. and -2.5 m.y., conifers, except for Cryptomeria types, were prominent, Quercus continued to decline, and other broad-leaf trees were minor. Over the last 2 Ma, the very large and frequent changes in forest composition inferred from pollen in the Sea of Japan correspond to forest dynamics inferred from changes in pollen and floral assemblages throughout Japan. Given present vegetation/climate relationships, broad trends in Neogene climate inferred from these preliminary pollen data include decreasing temperatures, increasing seasonality in temperatures and precipitation, and increasing amplitude and frequency of climatic change. Two significant events, centered at ~9 m.y. and ~4 m.y., punctuate the gradual deterioration of the equable warm, humid subtropical/warm temperate late Miocene and early Pliocene climates. The first indication of cold-temperate conditions comparable to those of Pleistocene glacial intervals occurs ~3 m.y. Subsequently, regional climates oscillated rapidly between temperate and cold-temperate regimes that supported conifer and mixed broad-leaf forests; however, climatic extremes were apparently never great enough to displace warm-temperate and temperate forests from Honshu nor to produce arctic climates on the west coast of Japan.

Pollen profile and age determination for sediment core M72/5_619-1 (22-GC3)

Sediments from the Black Sea, a region historically dominated by forests and steppe landscapes, are a valuable source of detailed information on the changes in regional terrestrial and aquatic environments at decadal to millennial scales. Here we present multi-proxy environmental records (pollen, dinoflagellate cysts, Ca, Ti and oxygen isotope data) from the uppermost 305 cm of the core 22-GC3 (42°13.53' N, 36°29.55' E) collected from a water depth of 838 m in the southern part of the Black Sea in 2007. The records span the last ~ 18 kyr (all ages are given in cal kyr BP). The pollen data reveal the dominance of the Artemisia-steppe in the region, suggesting rather dry/cold environments ~ 18-14.5 kyr BP. Warming/humidity increase during melt-water pulses (~ 16.1-14.5 kyr BP), indicated by d18O records from the 22-GC3 core sediment and from the Sofular Cave stalagmite, is expressed in more negative d13C values from the Sofular Cave, usually interpreted as the spreading of C3 plants. The records representing the interstadial complex (~ 14.5-12.9 kyr BP) show an increase in temperature and moisture, indicated by forest development, increased primary productivity and reduced surface run-off, whereas the switch from primary terrigenous to primary authigenic Ca origin occurs ~ 500 yr later. The Younger Dryas cooling is clearly demonstrated by more negative d13C values from the Sofular Cave and a reduction of pines. The early Holocene (11.7-8.5 kyr BP) interval reveals relatively dry conditions compared to the mostly moist and warm middle Holocene (8.5-5 kyr BP), which is characterized by the establishment of the species-rich warm mixed and temperate deciduous forests in the low elevation belt, temperate deciduous beech-hornbeam forests in the middle and cool conifer forest in upper mountain belt. The border between the early and middle Holocene in the vegetation records coincides with the opening of the Mediterranean corridor at ~ 8.3 kyr BP, as indicated by a marked change in the dinocyst assemblages and in the sediment lithology. Changes in the pollen assemblages indicate a reduction in forest cover after ~ 5 kyr BP, which was likely caused by increased anthropogenic pressure on the regional vegetation.

(Table T1) Kerogen and organic constituents in ODP Hole 183-1138A, Kerguelen Plateau

Mid-Cretaceous sediments recovered during Ocean Drilling Program Leg 183 (Cores 183-1138A-69R to 73R) on the central Kerguelen Plateau have been analyzed palynologically and paleobotanically to determine the age of the strata and to reconstruct vegetational development and paleoecology. The lower strata (Cores 183-1138A-71R to 73R), a dark, organic-rich silty claystone with many wood fragments and fern remains (sedimentary Unit VI), certainly of terrestrial origin, directly overlies the volcanic basement, which is dated as latest Albian (~95 to 103 Ma) to earliest Cenomanian. The age of the terrestrial strata can be determined by sporomorphs as late Albian to earliest Cenomanian as well. This shows that parts of the central Kerguelen Plateau must have been subaerial at least until the late Albian and were covered with a diverse high-latitude flora, probably dense conifer forest with various fern taxa in the undergrowth. Early angiosperms are also present. The vegetational character represented in Unit VI did not change significantly through time. However, varying percentages of several sporomorph groups seem to show recurring abundance variations, which might possibly be cyclic, caused by Milankovitch-type cyclic events. Cores 183-1138A-67R through 69R, of open marine origin, contain medium- to high-diversity dinocyst assemblages. Based on previous stratigraphic zonation schemes, the ages of these strata range within the Heterosphaeridium Superzone, from the Palaeohystrichophora infusorioides Zone to the Conosphaeridium striatoconus Zone, which correlates to the latest Cenomanian to Coniacian.

Mortality and climatic/environmental correlates of Widdringtonia cedargensis trees in the Cederberg, South Africa

Background: Conifer populations appear disproportionately threatened by global change. Most examples are, however, drawn from the northern hemisphere and long-term rates of population decline are not well documented as historical data are often lacking. We use a large and long-term (1931-2013) repeat photography dataset together with environmental data and fire records to account for the decline of the critically endangered Widdringtonia cedarbergensis. Eighty-seven historical and repeat photo-pairs were analysed to establish 20th century changes in W. cedarbergensis demography. A generalized linear mixed-effects model was fitted to determine the relative importance of environmental factors and fire-return interval on mortality for the species. Results: From an initial total of 1313 live trees in historical photographs, 74% had died and only 44 (3.4%) had recruited in the repeat photographs, leaving 387 live individuals. Juveniles (mature adults) had decreased (increased) from 27% (73%) to 8% (92%) over the intervening period. Our model demonstrates that mortality is related to greater fire frequency, higher temperatures, lower elevations, less rocky habitats and aspect (i.e. east-facing slopes had the least mortality). Conclusions: Our results show that W. cedarbergensis populations have declined significantly over the recorded period, with a pronounced decline in the last 30 years. Individuals that established in open habitats at lower, hotter elevations and experienced a greater fire frequency appear to be more vulnerable to mortality than individuals growing within protected, rocky environments at higher, cooler locations with less frequent fires. Climate models predict increasing temperatures for our study area (and likely increases in wildfires). If these predictions are realised, further declines in the species can be expected. Urgent management interventions, including seedling out-planting in fire-protected high elevation sites, reducing fire frequency in higher elevation populations, and assisted migration, should be considered.