The distribution of juvenile benthic invertebrates in an arctic glacial fjord

The distribution of juvenile benthic invertebrates with respect to their abiotic environment was investigated at four locations along a gradient in Kongsfjorden (79°N, 12°E), Spitsbergen, in September 1997. Heavy discharge of inorganic sediments released by the three glaciers was expected to be one of the main structuring factors of benthic communities in the fjord. Juveniles of 76 species/families are identified. Total abundance increases with distance from the fjord head due to the dominance of opportunistic polychaetes, while diversity is highest close to the glacier. Juvenile polychaetes are the most abundant taxon over all stations (92%), followed by bivalves, crustaceans and others. Polychaetes also provide most species (55%) and are the dominant taxon at every station. Their abundance is highly correlated with total organic carbon, indicating that most of them are deposit feeders. Young crustaceans rapidly decrease towards the fjord mouth, probably due to stronger bottom currents. Even though young oligochaetes are exclusively found near the glacier, no species seem to be well adapted to the high sedimentation rate close to the glacier. Juvenile suspension-feeding bivalves seem to be less disturbed by glacial discharge but appear to be more vulnerable to currents on the more exposed sites. The importance of deposit feeders and carnivores increases towards the outer stations. Non-metric multidimensional scaling confirms the distinct gradient in community composition along the fjord. Coupling the biotic data to abiotic factors (depth, bottom-water salinity, bottom-water temperature, sediment grain size and sedimentation rate) using canonical correspondence analysis revealed that hydrographic factors are more responsible for the structuring of the benthic juvenile community at the shallow stations close to the glacier (except the station directly at the glacier). At the outer deeper stations, sediment grain size and related properties may play a more important role.     

Distribution of meiofauna in Kongsfjorden,Spitsbergen

Kongsfjorden, a glacial fjord, is a typical fjord in the Spitsbergen (Svalbard archipelago) in the Arctic. The study supports a hypothesis that meiofauna and macrofauna are affected by natural environmental disturbances. Therefore, meiofaunal and macrofaunal analyses can be used to assess the effects of natural environmental disturbances in similar fjords in the Spitsbergen. Inputs from tidal glaciers create steep environmental gradients in sedimentation and salinity along the fjord. The magnitude of the glacial outflow diminishes towards the outer part of the fjord. Glacial-related physical stress causes reduced abundance, biomass and diversity among the meiofaunal assemblages in the inner part of the fjord. Based on quantitative and qualitative analyses of the composition of collected samples, three groups of meiofauna have been distinguished: one outer basin association and two in the inner, glacial bay. The presented results demonstrate that both the meiofauna and the macrofauna are affected on a similar scale by natural environmental disturbances. Therefore, as for macrofauna, meiofaunal analysis can be used to assess the effect of natural environmental disturbances.     

Meiofaunal distribution in Hornsund fjord,Spitsbergen

The meiofaunal community structure at 32 stations in Hornsund fjord (77°N) was investigated, and results were compared with data from another Spitsbergen fjord, Kongsfjorden (79°N). Steep environmental gradients of sedimentation, organic matter content, and salinity from the inner to the outer basin of the fjord are present due to intensive glacial discharges of meltwater and ice. As the natural environmental disturbances were described for macrofauna benthic communities before, we aimed to check whether the same pattern occurs among meiofauna. A total of 12 higher meiofaunal taxa were recorded, with nematodes predominating at all stations. Non-parametric multivariate analyses demonstrated clear differences in meiofaunal abundance and composition between stations in the glacial bay and in the outer part of the fjord. Meiofaunal abundance increased with increasing distance from the source of disturbance, which in our study is tidal glaciers. Therefore, the current study demonstrates that the spatial structure of meiofauna is affected by the natural environmental disturbance, and analysis of meiofaunal assemblages can be used to assess the effect of such disturbances.     

Composition of bryozoan assemblages related to depth in Svalbard fjords and sounds

Svalbard bryozoan communities were investigated along a depth range from the surface to 296 m between the inner glacial fronts and fjord mouths during 2001 and 2002. The main study area was Kongsfjorden (79°N, 12°E). A total of 137 taxa of bryozoans were identified: 108 to species, 24 to genus, 3 to family, 1 to order and 1 to phylum level. Cluster and multidimensional scaling analyses revealed four distinct assemblages of bryozoans: shallow (0–40 m; 68 taxa), deep (40–296 m; 80 taxa), inner fjordic (three taxa) and an assemblage found on small stones in shallow waters (nine taxa). The inner fjordic assemblage was recorded from the front of tidal glaciers extending about 10 km out into the fjord. In terms of abundance, Celleporella hyalina Linnaeus dominated in shallow areas (18%), Hippothoa arctica Kluge (55%) in deep water, Alcyonidium disciforme Smitt (86%) proximate to glaciers fronts and Electra arctica Borg on small stones (98%). The species were classified according to their depth range as a stenobathic-shallow (46 taxa), stenobathic-deep (57 taxa) and eurybathic-generalist (21 taxa). Mean diversity measures did not show any significant differences between the shallow and deep communities. The bryozoan assemblages seem to be structured primarily by processes related to depth and sediment characteristics.     

Distribution patterns in the biomass of macrozoobenthic communities in Admiralty Bay (King George Island, South Shetlands, Antarctic)

The distribution patterns of macrozoobenthic communities as shown by the biomass of the higher taxa were investigated in Admiralty Bay. Material was collected at depths ranging from 4 to 500 m (102 quantitative samples), representing the full depth range of this basin and including areas characterized by the different levels of glacial influence. Five community groups were distinguished by multivariate analysis (clustering, MDS) based on the Bray--Curtis similarity index. In the Ezcurra Inlet, the area characterized by intensive glacial processes resulting in a high sedimentation rate, the distribution of biomass was not depth related, but was associated with the intensity of glacial disturbance along the axis of the fjord. The innermost part of this fjord, which receives strong outflows from glaciers, had a low biomass value (2.9 g/0.1 m²) and was dominated by polychaetes. The middle part of the fjord which is not so heavily disturbed had higher biomass values (46.1 g/0.1 m²) due to the dominance of bivalves. A clear depth gradient of biomass distribution was observed in bottom areas located far from glaciers, in the central basin of the bay, with three zones within the depth ranges 4–30, 40–380, and 400–500 m. The highest mean biomass values (92.5 g/0.1 m²) were found in the middle sublittoral (40–380 m) dominated by ascidians. In the shallow and deep assemblages, the biomass was lower (33.8 g/0.1 and 30.9 g/0.1 m² respectively), most probably associated with the ice disturbance in the shallows and lower food supply in the deepest part of the shelf.     

Decadal change in macrobenthic soft-bottom community structure in a high Arctic fjord (Kongsfjorden, Svalbard)

Marine benthic macrofauna communities are considered a good indicator of subtle environmental long-term changes in an ecosystem. In 1997/1998 and 2006, soft-bottom fauna of an Arctic glacial fjord Kongsfjorden was extensively sampled and major communities were identified along the fjord axis, which were related to the diminishing influence of glacial activity. Spatial patterns in community structure and species diversity were significantly different in the central basin of Kongsfjorden between periods while there was no change in the inner part of the fjord. In 1997/98, three faunal associations were distinguished with significant differences in species richness and diversity (H′) while in 2006 only two faunal associations were identified and there were no differences any more between the two formerly distinct associations in the central fjord. The increased input of Atlantic water due to a stronger West Spitsbergen Current may be the reason for unification of previous clear faunal division. The faunal association in the inner, well separated glacial part of the fjord, characterized by strong glacier influence, was protected from Atlantic water inflow and, hence, the macrobenthic fauna essentially remained unaffected. Reduced abundance of species typical for glacial bays in the central part of the fjord in 2006 may result from the decreasing effect of Blomstrandbreen glacier, strong increase of input of Atlantic water into the fjord and increased temperature of West Spitsbergen Current. Higher values of POC in 2006 than in 1998 are likely the effect of increased primary production resulting from warmer water temperatures.     

Sedimentation and composition of wall communities in Alaskan fjords

The effects of suspended sediments on the composition of wall communities in Alaskan fjords were investigated by quantitative assessment of underwater photo-quadrats. In fjords with actively retreating tidewater glaciers, suspended sediment levels were extremely high at the heads and were exponentially lower at the mouths. Fjords without glaciers had low suspended sediment levels throughout. The per cent cover and number of species were lowest where sedimentation was highest, at the heads of glacial fjords. Here the wall communities were dominated by a sparse cover of small serpulid worms. Richer communities comprising mostly algae, sponges, hydroids, bryozoans, tunicates and brachiopods were found in fjords without glaciers, and at the mouths of glacial fjords. There was a positive correlation between water clarity and community composition along a gradient from heads to the mouths of fjords with high suspended sediment levels. Accepted: 11 January 1999     

Macrobenthic species composition and diversity in the Godthaabsfjord system, SW Greenland

The southwest Greenland coast is made up of large and deep sill fjords. On the shelf, a number of shallow banks separated by deep troughs are located 20–50 km from the coast. We collected three 0.1-m² van Veen grabs at nine stations along a transect spanning from the inner Godthaabsfjord influenced by glaciers, across the shallow Fyllas Bank and out to the slope of the continental shelf at approximately 1,000 m depth. Along this transect, we explored patterns of macrobenthic diversity, species composition, abundance and biomass. The sampled stations were very different in terms of environmental variables, resulting in large differences in species composition primarily related to differences in depth, silt–clay fraction and chl a content of the sediment (BIO-ENV analysis). Habitat differences also reduced species spatial ranges and the majority of species were found at only one (49%) or two (20%) stations and, consequently, species turnover or beta diversity was high and correlated to differences in depth, silt–clay fraction and median sediment grain size. Species richness and diversity were lowest in sites exposed to sediment disturbance: near the glaciers in the inner fjord (physical disturbance by mineral sedimentation) and at selected stations on the shelf (bioturbation by burrowing sand eel). Alpha diversity and richness were only weakly correlated to environmental parameters, indicating that alpha richness and diversity are influenced by several factors or that relationships are non-linear as was found for species richness and silt–clay fraction.     

Marine‐terminating glaciers sustain high productivity in Greenland fjords

Accelerated mass loss from the Greenland ice sheet leads to glacier retreat and an increasing input of glacial meltwater to the fjords and coastal waters around Greenland. These high latitude ecosystems are highly productive and sustain important fisheries, yet it remains uncertain how they will respond to future changes in the Arctic cryosphere. Here we show that marine‐terminating glaciers play a crucial role in sustaining high productivity of the fjord ecosystems. Hydrographic and biogeochemical data from two fjord systems adjacent to the Greenland ice sheet, suggest that marine ecosystem productivity is very differently regulated in fjords influenced by either land‐terminating or marine‐terminating glaciers. Rising subsurface meltwater plumes originating from marine‐terminating glaciers entrain large volumes of ambient deep water to the surface. The resulting upwelling of nutrient‐rich deep water sustains a high phytoplankton productivity throughout summer in the fjord with marine‐terminating glaciers. In contrast, the fjord with only land‐terminating glaciers lack this upwelling mechanism, and is characterized by lower productivity. Data on commercial halibut landings support that coastal regions influenced by large marine‐terminating glaciers have substantially higher marine productivity. These results suggest that a switch from marine‐terminating to land‐terminating glaciers can substantially alter the productivity in the coastal zone around Greenland with potentially large ecological and socio‐economic implications.     

Distribution patterns of polychaete fauna in an Arctic fjord (Hornsund, Spitsbergen)

Polychaetes are one of the most important groups of benthic organisms in marine ecosystems. They dominate on the Arctic shelf and play an important role in ecosystem functioning. This study focuses on the polychaete biodiversity and their distribution patterns in Hornsund, an open glacial fjord, in western Spitsbergen and provides important baseline data for future studies of temporal fluctuations in benthic fauna. The main aim of this study was to assess how the polychaete abundance, biomass, diversity, community structure, and function vary along the Hornsund fjord’s axis, in relation to the environmental factors. Eighty-eight polychaete taxa were identified; an average density was 457 ind. m^?2 ± 237.5 SD. Three assemblages were distinguished (INNER, MIDDLE, and OUTER) along the fjord axis, reflecting the intensity of glacial disturbance. A clear division between the highly disturbed inner part of the fjord (Brepollen) and the less impacted middle and outer parts was observed. Continuous gradients in abundance, biomass, and diversity were found with all those values diminishing toward the inner region of the fjord. The polychaete assemblages' indices were significantly correlated with bottom temperature, sediment characteristics (grain size), and distance to the glacier (longitude). No significant correlations were found with depth or total organic carbon content. Carnivore and motile surface deposit feeding polychaete species dominated in the areas close to the glaciers, while the OUTER community was dominated by carnivores and surface sessile and discretely motile species, and had more complex trophic structure, with multiple species representing different functional groups including carnivores, sessile, discretely motile, and motile surface deposit feeders and motile burrowers.     

Distinct or similar? Soft bottom polychaete diversity in Arctic and Antarctic glacial fjords

The main aim of this study was to compare the polychaete communities in two similar polar areas: an Arctic fjord, Hornsund (Svalbard) and an Antarctic fjord, Ezcurra Inlet (South Shetlands). This is the first attempt to compare Arctic and Antarctic diversity based on fully comparable datasets. Forty van Veen grab samples were collected in each fjord: twenty replicates were taken in each of two fjord areas characterized by a different level of glacial disturbance—in the inner (glacial bay) and outer (fjord mouth) region of both fjords, from depths of about 100 m in 2005 (Hornsund) and in 2007 (Ezcurra Inlet). In the glacial bays, species richness and diversity were significantly higher in Ezcurra Inlet than in Hornsund due to higher rate of glacial disturbance in the latter one. In the outer areas, species richness was similar in both fjords, although diversity values were higher in Ezcurra Inlet. Polychaete species richness in the habitats characterized by similar level of disturbance (outer areas of the fjords) was the same in both polar regions. At this small scale, where community drivers are very similar, the species richness seems to be independent from the local or regional species pool.     

Free-living nematodes and macrobenthos in a high-latitude glacial fjord

Kongsfjord is an open glacial fjord on the west coast of Svalbard, where the influence of the West Spitzbergen current ameliorates the effects of high latitude (79°N). The fjord is heavily influenced by glacial discharges of meltwater, ice and till, and related environmental gradients in sediments from the glaciers to the open sea include sediment deposition, organic content and disturbance. Other factors, such as the formation and break up of sea ice, also affect benthic communities. In this study spatial patterns in nematode and macrofaunal communities, in samples collected using box-corers and van Veen grabs during a cruise in September 1997, are described, compared and contrasted. Non-parametric multivariate analyses demonstrate that there were clear differences in community structure between stations in both macrofaunal and nematode assemblages. At stations where macrofauna were sampled using both box-cores and grabs there were also significant differences between samples collected by different methods, although there is evidence that these were influenced in part by slight differences in sampling location. Some evidence of disturbance to macrofaunal assemblages in the centre of the fjord is apparent. Macrofaunal community composition varied most closely with a combination of depth and sediment C : N ratio, whereas that of nematodes varied most closely with C : N alone. Proportions of feeding groups of nematodes showed little variation along the fjord. There is no evidence of a specialised nematode assemblage inhabiting the part of the fjord subject to the heaviest deposition of sediment. The taxonomic distinctness of nematodes decreased with increasing distance from the source of disturbance. This is in contrast to studies showing that the taxonomic distinctness of nematodes tends to decrease with increasing anthropogenic stress.     

Megafaunal Communities in Rapidly Warming Fjords along the West Antarctic Peninsula: Hotspots of Abundance and Beta Diversity

Glacio-marine fjords occur widely at high latitudes and have been extensively studied in the Arctic, where heavy meltwater inputs and sedimentation yield low benthic faunal abundance and biodiversity in inner-middle fjords. Fjord benthic ecosystems remain poorly studied in the subpolar Antarctic, including those in extensive fjords along the West Antarctic Peninsula (WAP). Here we test ecosystem predictions from Arctic fjords on three subpolar, glacio-marine fjords along the WAP. With seafloor photographic surveys we evaluate benthic megafaunal abundance, community structure, and species diversity, as well as the abundance of demersal nekton and macroalgal detritus, in soft-sediment basins of Andvord, Flandres and Barilari Bays at depths of 436–725 m. We then contrast these fjord sites with three open shelf stations of similar depths. Contrary to Arctic predictions, WAP fjord basins exhibited 3 to 38-fold greater benthic megafaunal abundance than the open shelf, and local species diversity and trophic complexity remained high from outer to inner fjord basins. Furthermore, WAP fjords contained distinct species composition, substantially contributing to beta and gamma diversity at 400–700 m depths along the WAP. The abundance of demersal nekton and macroalgal detritus was also substantially higher in WAP fjords compared to the open shelf. We conclude that WAP fjords are important hotspots of benthic abundance and biodiversity as a consequence of weak meltwater influences, low sedimentation disturbance, and high, varied food inputs. We postulate that WAP fjords differ markedly from their Arctic counterparts because they are in earlier stages of climate warming, and that rapid warming along the WAP will increase meltwater and sediment inputs, deleteriously impacting these biodiversity hotspots. Because WAP fjords also provide important habitat and foraging areas for Antarctic krill and baleen whales, there is an urgent need to develop better understanding of the structure, dynamics and climate-sensitivity of WAP subpolar fjord ecosystems.     

Influence of Deglaciation on Microbial Communities in Marine Sediments Off the Coast of Svalbard,Arctic Circle

Increases in global temperatures have been shown to enhance glacier melting in the Arctic region. Here, we have evaluated the effects of meltwater runoff on the microbial communities of coastal marine sediment located along a transect of Temelfjorden, in Svalbard. As close to the glacier front, the sediment properties were clearly influenced by deglaciation. Denaturing gradient gel electrophoresis profiles showed that the sediment microbial communities of the stations of glacier front (stations 188–178) were distinguishable from that of outer fjord region (station 176). Canonical correspondence analysis indicated that total carbon and calcium carbonate in sediment and chlorophyll a in bottom water were key factors driving the change of microbial communities. Analysis of 16S rRNA gene clone libraries suggested that microbial diversity was higher within the glacier–proximal zone (station 188) directly affected by the runoffs than in the outer fjord region. While the crenarchaeotal group I.1a dominated at station 176 (62%), Marine Benthic Group-B and other Crenarchaeota groups were proportionally abundant. With regard to the bacterial community, alpha-Proteobacteria and Flavobacteria lineages prevailed (60%) at station 188, whereas delta-Proteobacteria (largely sulfate-reducers) predominated (32%) at station 176. Considering no clone sequences related to sulfate-reducers, station 188 may be more oxic compared to station 176. The distance-wise compositional variation in the microbial communities is attributable to their adaptations to the sediment environments which are differentially affected by melting glaciers.     

Disappearing Kilimanjaro snow—Are we the last generation to explore equatorial glacier biodiversity?

Glaciation accompanied our human ancestors in Africa throughout the Pleistocene. Regrettably, equatorial glaciers and snow are disappearing rapidly, and we are likely the last generation who will get to know these peculiar places. Despite the permanently harsh conditions of glacier/snow habitats, they support a remarkable diversity of life ranging from bacteria to animals. Numerous papers have been devoted to microbial communities and unique animals on polar glaciers and high mountains, but only two reports relate to glacial biodiversity in equatorial regions, which are destined to melt completely within the next few decades. Equatorial glaciers constitute “cold islands” in tropics, and discovering their diversity might shed light on the biogeography, dispersal, and history of psychrophiles. Thus, an opportunity to protect biota of equatorial glaciers hinges on ex situ conservation. It is timely and crucial that we should investigate the glacial biodiversity of the few remaining equatorial glaciers.     

Intertidal and subtidal soft-bottom macro- and meiofauna of the Kongsfjord (Spitsbergen)

The intertidal and subtidal soft bottom macro- and meiofauna of a glacier fjord on Spitsbergen was studied after complete ice melt in June 2003. The abundances of the benthic fauna were within the range reported from estuaries and similar intertidal areas of boreal regions. The high proportion of juveniles in the eulittoral zone indicated larval recruitment from subtidal areas. The macrobenthic fauna can be divided into an intertidal and a subtidal community, both being numerically dominated by annelids. Deposit feeders were numerically predominant in intertidal sites, whereas suspension feeders were most abundant in the subtidal area. Among the meiofauna, only the benthic copepods were identified to species, revealing ecological adaptations typical for intertidal species elsewhere.     

Loss of small glaciers will diminish beta diversity in Pyrenean streams at two levels of biological organization

Aim Small (< 1 km²) alpine glaciers are likely to disappear in this century, resulting in decreased regional habitat heterogeneity in associated streams. Both heterogeneity within and spatial isolation among glacier‐influenced streams can enhance beta diversity of stream‐dwelling organisms. We measured beta at both community and population‐genetic levels within and among streams currently influenced by small Pyrenean glaciers. We aimed to evaluate whether patterns are analogous between the two levels, to apply various approaches for characterizing beta, and to infer the outcome of future glacier loss on regional biodiversity. Location Four glacier‐fed basins in the Parc National des Pyrénées, France. Methods We classified each of 18 stream reaches across the basins into either high‐, mid‐ or low‐‘glaciality’ (glacial influence) groups according to four physicochemical characteristics. At each reach, we collected macroinvertebrate communities and evaluated mitochondrial DNA haplotypes for 11–13 individuals of Baetis alpinus Pictet. Using taxa/haplotypes as basic units, we evaluated community and population‐genetic beta diversity simultaneously. We measured beta diversity in three major ways: as multivariate (Sørensen's dissimilarity, Jost D) and ‘classical’ (gamma/alpha) variation to compare among glaciality groups, and as turnover along the glaciality gradient within each basin. Results For most approaches at both organizational levels, beta was greatest among high‐glaciality reaches, absolute values of variation of beta in high‐glaciality streams were strikingly similar between levels, and the steepest turnover within basins occurred between high‐ and mid‐glaciality reaches. Therefore, high‐glaciality reaches contained assemblages and populations that were unique both within that stream type (among basins) and compared with other stream types within basins. Main conclusions Parallel beta diversity patterns at population‐genetic and community levels suggested that environmental drivers influence these levels analogously. Extreme conditions (e.g. low temperature, high instability, isolation) in high‐glaciality streams probably enhance beta at both levels. Stream beta diversity is likely to decrease substantially with continued glacial reduction in this system.     

Direct and indirect effects of glaciers on aquatic biodiversity in high Andean peatlands

The rapid melting of glacier cover is one of the most obvious impacts of climate change on alpine ecosystems and biodiversity. Our understanding of the impact of a decrease in glacier runoff on aquatic biodiversity is currently based on the ‘glacier‐heterogeneity‐diversity’ paradigm, according to which there is high α‐diversity at intermediate levels of glacial influence due to the high degree of environmental heterogeneity caused by glacier water. This α‐diversity pattern generates high levels of between‐site aquatic community variation (high β diversity) and increases regional diversity (γ‐diversity). There is a rich conceptual background in favor of this paradigm, but empirical data supporting it are scarce. We investigated this paradigm by analyzing the different diversity patterns (α, β and γ‐diversity) of four aquatic groups (zooplankton, macroinvertebrates, algae and macrophytes) living in high‐elevation peatlands (>4500 m above sea level). We sampled 200 pools from 20 peatlands along a glacier gradient in the Cordillera Real of Bolivia. We performed structural equation modeling (SEM) to analyze the potential mechanisms underlying the observed diversity patterns. Intermediate levels of glacial influence (15–20% cover) resulted in high heterogeneity, but α‐diversity responded to glacial influence only for the zooplankton group (Cladocera). Our SEM analysis did not identify environmental heterogeneity as a significant variable explaining the relationship between glacier and α‐diversity. Peatland area had a strong positive effect on heterogeneity and diversity. β‐diversity was significantly associated with glacier gradient, and 12.9% of the total regional diversity (γ‐diversity) was restricted to peatlands with a high degree of glacial influence. These species might be lost in a context of glacial retreat. These findings provide new insight into the potential effects of glacial retreat on the aquatic environment and biodiversity in the peatlands of the tropical Andes.     

Keep your feet warm? A cryptic refugium of trees linked to a geothermal spring in an ocean of glaciers

Up to now, the most widely accepted idea of the periglacial environment is that of treeless ecosystems such as the arctic or the alpine tundra, also called the tabula rasa paradigm. However, several palaeoecological studies have recently challenged this idea, that is, treeless environments in periglacial areas where all organisms would have been exterminated near the glacier formed during the Last Glacial Maximum, notably in the Scandinavian mountains. In the Alps, the issue of glacial refugia of trees remains unanswered. Advances in glacier reconstructions show that ice domes did not cover all upper massifs, but glaciers filled valleys. Here, we used fossils of plant and malacofauna from a travertine formation located in a high mountain region to demonstrate that trees (Pinus, Betula) grew with grasses during the Lateglacial‐Holocene transition, while the glacier fronts were 200–300 m lower. The geothermal travertine started to accumulate more than 14,500 years ago, but became progressively more meteogene about 11,500 years ago due to a change in groundwater circulation. With trees, land snails (gastropods) associated to woody or open habitats and aquatic mollusc were also present at the onset of the current interglacial, namely the Holocene. The geothermal spring, due to warm water and soil, probably favoured woody glacial ecosystems. This new finding of early tree growth, combined with other scattered proofs of the tree presence before 11,000 years ago in the western Alps, changes our view of the tree distribution in periglacial environments, supporting the notion of tree refugia on nunataks in an ocean of glaciers. Therefore, the tabula rasa paradigm must be revisited because it has important consequences on the global changes, including postglacial plant migrations and biogeochemical cycles.     

Microphytic standing stocks at Kerguelen Islands (Subantarctic,Indian Ocean), annual variations in relation to environmental factors: II,intertidal and subtidal microphytobenthos

The microphytobenthic standing crop in marine coastal sediments from Kerguelen main island (from surface to 8 cm deep) varied in relation to the tidal position, grain size, and shelter of sediments. A time-series analysis was performed at three distinct sites: a sheltered fjord (PRAY), a moderately exposed embayment (PAF) and a deep fjord with higher open-ocean influence (PN). Temporal variations on all studied parameters were observed on intertidal, as well as on subtidal sediments, but with a great range in variations and pattern. Compared to phytoplankton blooms at the same location and stations, the microphytobenthos productive periods were not so marked, especially on intertidal sediments. Nevertheless, high productivity periods were in most cases linked to austral spring and summer (up to 30–50 g Chl a g^–1 dw, in intertidal sheltered sands; >170 g Chl a g^–1 dw, in sheltered subtidal muds). On subtidal muddy sites, the high phaeopigments concentrations (up to 195 g Phaeo a g^–1 dw at PRAY site) were attributed to kelp and epiphyte degradation and sedimentation, and also to macrofauna trophic influence, whereas at Portes-Noires fjord (PN) a Phaeophytine a late spring increase in 1991 was suspected to be of plankton origin. The PN site exhibited a lower productivity than at the other two sites, regardless of the tidal position of the sediment, due to a delayed annual cycle in surficial temperature and standing crop. Compared to the phytoplankton blooms, the microphytobenthos productivity at Kerguelen main island was less restricted in time and may occur throughout the year, not just to the benefit of the benthos food-chain, but also to the pelagic one via resuspension, as well as to filter feeders (i.e. mussels).