Plant species, biomass and environmental characteristics of relevés along the North America Arctic bioclimate gradient

Question: How do interactions between the physical environment and biotic properties of vegetation influence the formation of small patterned-ground features along the Arctic bioclimate gradient? Location: At 68° to 78°N: six locations along the Dalton Highway in arctic Alaska and three in Canada (Banks Island, Prince Patrick Island and Ellef Ringnes Island). Methods: We analysed floristic and structural vegetation, biomass and abiotic data (soil chemical and physical parameters, the n-factor [a soil thermal index] and spectral information [NDVI, LAI]) on 147 microhabitat releves of zonalpatterned-ground features. Using mapping, table analysis (JUICE) and ordination techniques (NMDS). Results: Table analysis using JUICE and the phi-coefficient to identify diagnostic species revealed clear groups of diagnostic plant taxa in four of the five zonal vegetation complexes. Plant communities and zonal complexes were generally well separated in the NMDS ordination. The Alaska and Canada communities were spatially separated in the ordination because of different glacial histories and location in separate floristic provinces, but there was no single controlling environmental gradient. Vegetation structure, particularly that of bryophytes and total biomass, strongly affected thermal properties of the soils. Patterned-ground complexes with the largest thermal differential between the patterned-ground features and the surrounding vegetation exhibited the clearest patterned-ground morphologies.

(Table 1) Dinoflagellate cysts and acritarchs in upper Pliocene and Pleistocene sediments from ODP Hole 151-911A

Pliocene and Pleistocene sediments from Ocean Drilling Program Leg 151, Hole 911 A, drilled on the innermost Yermak Plateau (Eastern Arctic Ocean), were studied for their dinoflagellate cyst content. Three assemblage zones were tentatively defined, characterized by the predominance of few species. The composition of the assemblages changed markedly, even within single assemblage zones, during the last 2.6 to 2.8 m.y., reflecting the variable influence of warmer water from the Norwegian Sea, fluctuations in the influence of cold polar water masses, and the extent of sea-ice cover. Polar to subpolar surface water masses prevailed on the Yermak Plateau during the late Pliocene, when the eastern Arctic Ocean was probably isolated from the Norwegian-Greenland Sea. Intrusions of warmer water are recorded since the latest Pliocene, alternating with colder periods and a prolonged seasonal sea-ice cover. The composition of the dinoflagellate cyst assemblages has also changed considerably since the middle Pleistocene, reflecting the establishment of stronger fluctuations in surface water mass conditions than before at Yermak Plateau.

Dinoflagellate cyst records from the East Greenland continental margin

Late Weichselian and Holocene dinoflagellate cyst assemblages have been investigated at two stations situated close to the modern Polar Front at the continental margin oft East Greenland. Both the concentrations of dinoflagelate cysts and the assemblage composition reflect changes in the surface water conditions, occurring in distinct steps during the past 15,000 years. Low concentrations of dinoflagellate cysts during Termination Ia suggest harsh environmental conditions, most probably caused by an extensive sea-ice cover and/or a high influx of low salinity meltwater. A surface water warming was recorded from 13,000 - 12,000 years BP, related to the inflow of warmer water trom the North Atlantic into the western Norwegian-Greenland Sea. The interval between Terminations la and Ib was characterized by a strong seasonality with an extensive sea-ice cover in winter and relatively warm surface waters in summer. At the transition to the Holocene, a reorganisation of the hydrography resulted in surface water conditions characteristic for the Holocene with three well-defined major water masses and oceanographic fronts The modern water mass conditions at both stations were established at the end of Termination Ib, around 6,400 to 6,800 years BP. In general, the influence of colder surface waters was more pronounced at the location off Scoresby Sund throughout the Holocene. Arctic water had the strongest influence at both stations in the middle Holocene. A progressive cooling with an increase in sea-ice cover is time-transgressivelyrecorded at both stations during the Holocene, indicating that the Polar Front moved to its present position or that branches of the zonal currents expanded from the East Greenland shell eastward during tlie last 3,000 years.

Dinoflagellate cysts analyses from sediment cores of the Po River discharge plume over the last two centuries

To obtain insight into the natural and/or human-induced changes in the trophic state of the distal portion of the Po River discharge plume over the last two centuries, high temporal resolution dinoflagellate cyst records were established at three sites. Cyst production rates appear to reflect the natural variability in the river's discharge, whereas cyst associations reflect the trophic state of the upper waters, which in turn can be related to agricultural development. The increased abundances of Lingulodinium machaerophorum and Stelladinium stellatum found as early as 1890 and 1920 correspond to the beginning of the industrial revolution in Italy and the first chemical production and dispersion of ammonia throughout Europe. After 1955, the increased abundances of these species and of Polykrikos schwartzii, Brigantedinium spp. and Pentapharsodinium dalei correspond to agriculturally induced alterations of the hypertrophic conditions. A slight improvement in water quality can be observed from 1987 onward.

Age model, radiocarbon dates and dinoflagellate cyst measurements from sediment core DA06-139G

A marine sediment core from Vaigat in Disko Bugt, West Greenland, has been analysed in terms of lithology, dinoflagellate cysts and foraminifera in order to evaluate the influence of oceanographic variability on West Greenland glacier stability. The data show that during the past 5200 years the Atlantic foraminiferal abundance in the subsurface waters of the West Greenland Current (WGC) episodically increased, indicating periods of increases in the inflow of subsurface warm Atlantic water at 2000 - 1500 cal. yr BP and 1300 cal. yr BP as well as periods of less pronounced increased bottom-water temperatures around 4700 - 4000 cal. yr BP, 3100 - 2800, 2600, 1000 - 800, 500 - 400, and at 200 cal. yr. The sedimentological and dinoflagellate cyst data indicate that these episodes with enhanced advection of Irminger Sea-derived waters are accompanied by increased iceberg rafting, which we link to increased iceberg calving in relation to destabilization of the Jakobshavn Isbrae. The long-term trend in the data documents the end of a late-Holocene Thermal Maximum between 5200 and 4300 cal. yr BP and a final onset of the Neoglaciation at 3500 cal. yr BP. Increased responses of the iceberg rafting after 3500 cal. yr BP, reflects a westward/seaward advance of the glacier margin in relation to onset of Neoglaciation and a development of the glacier into a floating tongue after 2000 cal. yr BP. A comparison of our record with a record from the eastern North Atlantic indicates that a NAO-like anomaly pattern between subsurface waters in West Greenland and atmospheric temperature in the Eastern North Atlantic may have been operating during most of the late Holocene. However, during the past 1000 years the NAO signal may have weakened as some other mode of climate variability overprints the anti-phase climate signal in this region.

Dinoflagellate cyst species composition of recent surface samples (Appendix)

Dinoflagellate cysts and other organic-walled microfossils have been studied in recent surface sediments from the entire Norwegian-Greenland Sea. More than 30 taxa have been recognized, of which only few show a distinct distribution pattern, and allow description of four assemblages. The occurrence of most taxa is related to the relatively warmer waters of the Norwegian Sea. Algidaspaeridium? minutum s.1., Brigantedinium simplex and Impagidinium? pallidum are the only species showing a preference for colder water masses. Two species, I.? pallidum and Nematosphaeropsis labyrinthus are mainly restricted to the oceanic environment, whereas the other species have also been reported from neritic environments in previous studies. Due to the limited knowledge of the ecological and sedimentological factors influencing the occurrence of dinoflagellate cysts in oceanic environments, their distribution in recent sediments can be only related to surface water masses in a broad sense. Although the distribution of assemblages correlates with specific surface water masses, comparison with assemblages recovered from sediment traps deployed basinwide in the Norwegian-Greenland Sea (Dale and Dale, 1992) revealed some major discrepancies in species composition and percentage abundances. The differences cannot be explained with certainty at the moment, although there is some evidence that transport of dinoflagellate cysts and other fossilizable microplankton in water masses by currents, in sea-ice and sediments may modify the assemblages found in recent oceanic surface sediments from the Norwegian-Greenland Sea.

Geographic distribution of dinoflagellate cysts in surface sediments

Dinoflagellate cysts are useful for reconstructing upper water conditions. For adequate reconstructions detailed information is required about the relationship between modern day environmental conditions and the geographic distribution of cysts in sediments. This Atlas summarises the modern global distribution of 71 organicwalled dinoflagellate cyst species. The synthesis is based on the integration of literature sources together with data of 2405 globally distributed surface sediment samples that have been preparedwith a comparable methodology and taxonomy. The distribution patterns of individual cyst species are being comparedwith environmental factors that are knownto influence dinoflagellate growth, gamete production, encystment, excystment and preservation of their organic-walled cysts: surface water temperature, salinity, nitrate, phosphate, chlorophyll-a concentrations and bottom water oxygen concentrations. Graphs are provided for every species depicting the relationship between seasonal and annual variations of these parameters and the relative abundance of the species. Results have been compared with previously published records; an overview of the ecological significance as well as information about the seasonal production of each individual species is presented. The relationship between the cyst distribution and variation in the aforementioned environmental parameters was analysed by performing a canonical correspondence analysis. All tested variables showed a positive relationship on the 99% confidence level. Sea-surface temperature represents the parameter corresponding to the largest amount of variance within the dataset (40%) followed by nitrate, salinity, phosphate and bottom-water oxygen concentration, which correspond to 34%, 33%, 25% and 24% of the variance, respectively. Characterisations of selected environments as well as a discussion about how these factors could have influenced the final cyst yield in sediments are included.

Dinoflagellate cyst accumulation rates of ODP Hole 146-893A from Santa Barbara Basin

The dinoflagellate cyst record from Ocean Drilling Program Hole 893A, Santa Barbara Basin, southern California, is examined at millennial-scale resolution for the past 40 kyr. Changes in cyst abundance, composition of cyst assemblages, and their diversity reflect major shifts in climate and ocean circulation in the region over this time interval. Throughout the sequence, dinoflagellate cyst assemblages are dominated by heterotrophic dinoflagellates. Brigantedinium spp. and other upwelling-related taxa such as Echinidinium and Protoperidinium americanum are abundant, indicating the continued influence of coastal upwelling on the basin during the late Quaternary. A significant increase in cyst accumulation rates is seen during the Holocene and, to a lesser extent, during shorter warming events such as Bolling/Allerod and Dansgaard-Oeschger interstadials, implying enhanced marine productivity during these periods. Cyst diversity is high during the Holocene. An increase in abundance of cysts produced by autotrophic dinoflagellates in the late Holocene suggests enhanced input of warm, nutrient-rich waters. In contrast, cyst assemblages from the Last Glacial Maximum exhibit a relatively low diversity and an increase in the cysts of heterotrophic dinoflagellates, in particular Selenopemphix nephroides. The presence of this taxon in association with Brigantedinium spp. implies substantial cooling of surface waters in the Santa Barbara Basin at that time.

Dinocyst assemblages of site HM52-43

Past sea-surface conditions over the northern North Atlantic during the last glacial maximum were examined from the study of 61 deep-sea cores. The last glacial maximum time slice studied here corresponds to an interval between Heinrich layers H2 and H1, and spanning about 20-16 ka on a 14C time scale. Transfer functions based on dinocyst assemblages were used to reconstruct sea-surface temperature, salinity, and sea-ice cover. The results illustrate extensive sea-ice cover along the eastern Canadian margins and sea-ice spreading, only during winter, over most of the northern North Atlantic. On the whole, much colder winter prevailed, despite relatively mild conditions in August (10-15°C at most offshore sites), thus suggesting a larger seasonal contrast of temperatures than today. Lower salinity than at present is reconstructed, especially along the eastern Canadian and Scandinavian margins, likely because of meltwater supply from the surrounding ice sheets. These reconstructions contrast with those established by CLIMAP on the basis of planktonic foraminifera. These differences are discussed with reference to the stratigraphical frame of the last glacial maximum, which was not the coldest phase of the last glacial stage. The respective significance of dinocyst and foraminifer records is also examined in terms of the thermohaline characteristics of surface waters and the vertical structure of upper water masses, which was apparently much more stratified than at present in the northern North Atlantic, thus preventing deep-water formation.

Relative abundance of dinoflagellate cysts in surface sediments and dinoflagellate counts from three sediment cores from the Atlantic sector of the Southern Ocean

In this study we investigate the potential of organic-walled dinoflagellate cysts (dinocysts) as tools for quantifying past sea-surface temperatures (SST) in the Southern Ocean. For this purpose, a dinocyst reference dataset has been formed, based on 138 surface sediment samples from different circum-Antarctic environments. The dinocyst assemblages of these samples are composed of phototrophic (gonyaulacoid) and heterotrophic (protoperidinioid) species that provide a broad spectrum of palaeoenvironmental information. The relationship between the environmental parameters in the upper water column and the dinocyst distribution patterns of individual species has been established using the statistical method of Canonical Correspondence Analysis (CCA). Among the variables tested, summer SST appeared to correspond to the maximum variance represented in the dataset. To establish quantitative summer SST reconstructions, a Modern Analogue Technique (MAT) has been performed on data from three Late Quaternary dinocyst records recovered from locations adjacent to prominent oceanic fronts in the Atlantic sector of the Southern Ocean. These dinocyst time series exhibit periodic changes in the dinocyst assemblage during the last two glacial/interglacial-cycles. During glacial conditions the relative abundance of protoperidinioid cysts was highest, whereas interglacial conditions are characterised by generally lower cyst concentrations and increased relative abundance of gonyaulacoid cysts. The MAT palaeotemperature estimates show trends in summer SST changes following the global oxygen isotope signal and a strong correlation with past temperatures of the last 140,000 years based on other proxies. However, by comparing the dinocyst results to quantitative estimates of summer SSTs based on diatoms, radiolarians and foraminifer-derived stable isotope records it can be shown that in several core intervals the dinocyst-based summer SSTs appeared to be extremely high. In these intervals the dinocyst record seems to be highly influenced by selective degradation, leading to unusual temperature ranges and to unrealistic palaeotemperatures. We used the selective degradation index (kt-index) to determine those intervals that have been biased by selective degradation in order to correct the palaeotemperature estimates. We show that after correction the dinocyst based SSTs correspond reasonably well with other palaeotemperature estimates for this region, supporting the great potential of dinoflagellate cysts as a basis for quantitative palaeoenvironmental studies.

Morphology of Cycladophora davisiana davisiana from the Atlantic Ocean

In recent years, temporal fluctuations in the abundance of C. d. davisiana have been used frequently as a highresolution stratigraphic and paleoenvironmental tool. The modern ecology and morphologic variation (temporal and geographic) of this radiolarian species is evaluated to ascertain its potential stratigraphic and paleoenvironmental significance. Statistics were obtained on the width and height of all C. d. davisiana segments from Pleistocene populations of differing ages from the Northern Hemisphere (Labrador Sea and Iceland-Faeroe Ridge) and Southern Hemisphere (Namibian shelf and Meteor Rise). Results reveal that segment height variations between and within populations are more conservative than segment width. The mean sizes of the thorax and first abdominal segment have distinguishable differences between C. d. davisiana found in the North and South Atlantic. All populations have no significant difference in first abdominal segment width, however, mean heights of this segment differ greatly between populations of the North and South Atlantic. Second abdominal segment sizes show no clear population grouping. Size differences in post-cephalic segment size of these populations would appear to be related to some isolation of gene pools and possibly unknown paleoenvironmental factors. Temporal changes in the postcephalic size of C. d. davisiana may be used to: (1) identify temporally equivalent peaks in abundance of the species in a given region, (2) possibly evaluate the degree of mixing of water'masses between regions, and (3) trace the initial spread of the species from its area of origin. Cleve's 1887 plankton samples, between Greenland and Spitzsbergen, were studied and used in conjunction with other data to make the following conclusions on the modern ecology of C. d. davisiana in the Arctic and Greenland-Norwegian Seas. (1) It is presently absent in surface water plankton samples, (2) it currently lives at depths below 500 m, where it is rare, (3) it does not live in the upper 200 m under Arctic ice but is rare at greater depths, (4) it is absent in the upper 200 m near permanent Greenland Sea ice where normal oceanic salinity prevails, and (5) it is most common in deep marginal fjord environments which may serve as a refuge for the species during interglacial periods. In the Atlantic Ocean, the abundance of C. d. davisiana does not exceed 1% of the assemblage between the Subtropical Convergence of each hemisphere. In the Norwegian and Labrador Seas the species may occasionally be in the range of 1-5% of the modern radiolarian assemblage and never more than 5% in the southern high latitudes. Apparently only in the modern Sea of Okhotsk, does the species presently occur in high abundance. We concur with Morley and Hays (1983) that increased abundances are likely caused by the development of a strong low-salinity surface layer associated with seasonal sea ice melting and a strong temperature minimum above warmer and higher salinity intermediate waters. Similar conditions were frequent during the Pleistocene in the high latitudes and its modern scarcity outside the Sea of Okhotsk must be related to the absence of the presently unique conditions in the latter region.