The variable colours of the fiddler crab Uca vomeris and their relation to background and predation

Colour changes in fiddler crabs have long been noted, but a functional interpretation is still lacking. Here we report that neighbouring populations of Uca vomeris in Australia exhibit different degrees of carapace colours, which range from dull mottled to brilliant blue and white. We determined the spectral characteristics of the mud substratum and of the carapace colours of U. vomeris and found that the mottled colours of crabs are cryptic against this background, while display colours provide strong colour contrast for both birds and crabs, but luminance contrast only for a crab visual system. We tested whether crab populations may become cryptic under the influence of bird predation by counting birds overflying or feeding on differently coloured colonies. Colonies with cryptically coloured crabs indeed experience a much higher level of bird presence, compared to colourful colonies. We show in addition that colourful crab individuals subjected to dummy bird predation do change their body colouration over a matter of days. The crabs thus appear to modify their social signalling system depending on their assessment of predation risk.

(Appendix B) Dinoflagellate cysts in Southeast Pacific Ocean surface sediments

orty-eight surface sediment samples from the southeast (SE) Pacific (25-53°S) are investigated for the determination of the spatial distribution of organic-walled dinoflagellate cysts along the western South American continental margin. Fifty-five different taxa are recorded and reflect oceanic or coastal assemblages. The oceanic assemblages are characterised by low cyst concentrations and the dominance of autotrophs, while the coastal assemblages generally contain a higher number of cysts, which are mainly produced by heterotrophic species. Highest cyst concentrations are observed in the active upwelling system offshore Concepción (35-37°S). Brigantedinium spp., Echinidinium aculeatum, Echinidinium granulatum/delicatum and cysts of Protoperidinium americanum dominate assemblages related to upwelling. Echinidinium aculeatum appears to be the best indicator for the presence of all year round active upwelling cells. Other protoperidinioid cysts may also occur in high relative abundances in coastal regions outside active upwelling systems, if the availability of nutrients, co-responsible for the presence/absence of their main food sources such as diatoms and other protists, is sufficient. The importance of nutrient availability as a determining environmental variable influencing cyst signals on a regional scale (SE Pacific) is demonstrated through statistical analyses of the data. Because of the importance of nutrients, uncertainties about the outcomes of quantitative sea-surface temperature (SST) reconstructions (Modern Analogue Technique) based on dinoflagellate cysts may arise, since no interaction between different hydrographical variables is considered in this approach. The combination of the SE Pacific surface sample dataset with other published cyst data from the Southern Hemisphere resulted in a database which includes 350 samples: the 'SH350 database'. This database is used to test the accuracy of the quantitative reconstructions by calculating and comparing the estimated versus observed values for each site. An attempt to perform quantitative SST reconstructions on the last 25 cal ka of site ODP1233 (41°S; 74°27'W) is made and again stresses the importance of other environmental variables such as nutrient availability in determining the dinoflagellate cyst assemblages.

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.

Ostracode relative abundances and geochemistry in surface sediments from Lake Ohrid

Understanding the ecology of bioindicators such as ostracods is essential in order to reconstruct past environmental and climate change from analysis of fossil assemblages preserved in lake sediment cores. Knowledge of the ecology of ancient Lake Ohrid's ostracod fauna is very limited and open to debate. In advance of the Ohrid ICDP-Drilling project, which has potential to generate high-resolution long-term paleoenvironmental data of global importance in paleoclimate research, we sampled Lake Ohrid and a wide range of habitat types in its surroundings to assess 1) the composition of ostracod assemblages in lakes, springs, streams, and short-lived seasonal water bodies, 2) the geographical distribution of ostracods, and 3) the ecological characteristics of individual ostracod species. In total, 40 species were collected alive, and seven species were preserved as valves and empty carapaces. Of the 40 ostracod species, twelve were endemic to Lake Ohrid. The most common genus in the lake was Candona, represented by 13 living species, followed by Paralimnocythere, represented by five living species. The most frequent species was Cypria obliqua. Species with distinct distributions included Heterocypris incongruens, Candonopsis kingsleii, and Cypria lacustris. The most common species in shallow, flooded areas was H. incongruens, and the most prominent species in ditches was C. kingsleii. C. lacustris was widely distributed in channels, springs, lakes, and rivers. Statistical analyses were performed on a "Lake Ohrid" dataset, comprising the subset of samples from Lake Ohrid alone, and an "entire" dataset comprising all samples collected. The unweighted pair group mean average (UPGMA) clustering was mainly controlled by species-specific depth preferences. Canonical Correspondence Analysis (CCA) with forward selection identified water depth, water temperature, and pH as variables that best explained the ostracod distribution in Lake Ohrid. The lack of significance of conductivity and dissolved oxygen in CCA of Ohrid data highlight the uniformity across the lake of the well-mixed waters. In the entire area, CCA revealed that ostracod distribution was best explained by water depth, salinity, conductivity, pH, and dissolved oxygen. Salinity was probably selected by CCA due to the presence of Eucypris virens and Bradleystrandesia reticulata in short-lived seasonal water bodies. Water depth is an important, although indirect, influence on ostracod species distribution which is probably associated with other factors such as sediment texture and food supply. Some species appeared to be indicators for multiple environmental variables, such as lake level and water temperature.

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.

Dinoflagellate cysts in surface sediments

Surface sediments at 439 sites throughout the North Atlantic Ocean and adjacent seas have been analyzed for dinoflagellate cysts in order to establish a reference database from which paleoenvironmental transfer functions can be developed. Laboratory procedures and systematics were standardized in order to avoid bias introduced by the selective loss of taxa and to facilitate site to site comparison. 371 sites were retained to develop the database that includes 41 taxa, some of which were grouped using morphological and/or ecological criteria. 27 taxa were retained for statistical purposes. Distribution maps of these latter taxa have been plotted on the basis of their relative abundance. Principal component analyses were performed in order to describe the distribution of assemblages. The relation between the assemblages, as well as the relative abundance of individual taxa, and selected sea-surface parameters are illustrated. The parameters which were considered include temperature and salinity for winter (February) and summer (August) together with the duration of sea-ice cover. Transfer functions using the best analogue method have been tested with a view to reconstruct past sea-surface parameters. Validation procedures on this transfer function demonstrate that more than 95% of the reconstructions are included within the interannual variability of modern sea-surface conditions. Therefore, these transfer functions give accurate results and can be applied to reconstructing paleo-temperatures and -salinities from analogous assemblages in Quaternary sedimentary sequences. Protoperidinium stellatum (Wall in Wall & Dale, 1968) Head, comb. nov. (basionym = Peridinium stellatum) is proposed as new, and Algidasphaeridium? minutum var. cezare de Vernal et al., 1989 ex de Vernal et al. is newly validated.

Abundance of organic-walled dinoflagellate cysts in surface sediments from the Atlantic sector of the Southern Ocean (Appendix 2)

Thirty-two surface sediment samples from the Southern Ocean (eastern Atlantic sector), between the Subtropical Front and the Weddell Gyre, were investigated to provide information on the distribution of modern organic-walled dinoflagellate cysts in relation to the oceanic fronts of the Antarctic Circumpolar Current (ACC). A clearly distinguishable distribution pattern was observed in relation to the water masses and fronts of the ACC. The dinoflagellate cysts of species characteristic of open oceanic environments, such as Impagidinium species, are highly abundant around the Subtropical Front, whereas south of this front, cosmopolitan species such as Nematosphaeropsis labyrinthus and the cysts of Protoceratium reticulatum characterise the transition from subtropical to subantarctic surface waters. The subantarctic surface waters are dominated by the cysts of heterotrophic dinoflagellates, such as Protoperidinium spp. and Selenopemphix antarctica. The cysts of Protoperidinium spp. form the dominant part of the assemblages around the Antarctic Polar Front, whereas S. antarctica concentrations increase further to the south. The presence of S. antarctica in sediments of the Maud Rise, a region of seasonal sea-ice cover, reflects its tolerance for low temperatures and sea-ice cover. A previously undescribed species, Cryodinium meridianum gen. nov. sp. nov., has a restricted distribution pattern between the Antarctic Polar Front and the ACC-Weddell Gyre Boundary.

Micropaleontological investigation of sediment core Kronsmoor

Benthic foraminiferal and calcareous nannofossil assemblages, as well as stable isotope data from the Campanian/Maastrichtian boundary interval (~71.4 to ~70.7 Ma) of the Kronsmoor section (North German Basin), were investigated in order to characterize changes in surface-water productivity and oxygen content at the seafloor and their link to climatic and paleoceanographic changes. A nutrient index based on calcareous nannofossils is derived for the high-latitude, epicontinental North German Basin, reflecting changes in surface-water productivity. Oxygen isotopes of well-preserved planktic foraminiferal specimens of Heterohelix globulosa reflect warmer surface-water temperatures in the lower part of the studied succession and a cooling of up to 2°C (0.5 per mil) in the upper part (after 71.1 Ma). For the lower and warmer part of the investigated succession, benthic foraminiferal assemblages and the calcareous nannofossils indicate well-oxygenated bottom waters and low-surface water productivity. In contrast, the upper part of the succession is characterized by cooler conditions, lower oxygen content at the seafloor and increasing surface-water productivity. It is proposed that the cooling phase starting at 71.1 Ma was accompanied by increasing surface-water mixing caused by westerly winds. As a consequence of mixing, nutrients were advected from sub-surface waters into the mixed layer, resulting in increased surface-water productivity and enhanced organic matter flux to the seafloor. We hypothesize that global sea-level fall during the earliest Maastrichtian (~71.3 Ma), indicated by decreasing carbon isotope values, may have led to a weaker water mass exchange through narrower gateways between the Boreal Realm and the open North Atlantic and Tethys oceans. Both the weaker water mass exchange and enhanced surface-water productivity may have led to slightly less ventilated bottom waters of the upper part of the studied section. Our micro-paleontological and stable isotopic approach indicates short-term (<100 kyr) changes in oxygen consumption at the seafloor and surface-water productivity across the homogeneous Boreal White Chalk succession of the North German Basin.

Dinoflagellate cysts counts of sediment core MD95-2042

Dinoflagellate cysts were analysed from IMAGES core MD952042 (37°48?N; 10°01?W) retrieved from the Tagus Abyssal Plain. Previous results of stable isotope and magnetic susceptibility measurements as well as of planktonic foraminiferal temperature reconstruction from this core, suggest the occurrence of "Heinrich-like events" (i.e. large ice-sheet decay) during Marine Isotopic Stage 5 (MIS 5). Dinoflagellate assemblages of this time period have revealed six dinocyst events that are characterised by peaks in Bitectatodinium tepikiense percentages. These events occur synchronously with "Heinrich-like events" previously identified. They are coeval with major retreats of the forest on land, indicating, therefore, drastic changes in the regional climate. However, results from the Ice-Rafted Detritus (IRD) analysis of the >150 ?m lithic fraction shows that MIS 5 of MD952042 has only recorded one significant input of iceberg discharge, located at the MIS 6/MIS 5 transition. It seems therefore that it is the only event that could be called a "true Heinrich event".

Organic-walled dinoflagellate cysts and environmental analysis of surface sediments from the Northwest African upwelling area

In order to examine the spatial distribution of organic-walled dinoflagellate cysts (dinocysts) in recent sediments related to environmental conditions in the water column, thirty-two surface sediment samples from the NW African upwelling region (20-32°N) were investigated. Relative abundances of the dinocyst species show distinct regional differences allowing the separation of four hydrographic regimes. (1) In the area off Cape Ghir, which is characterized by most seasonal upwelling and river discharge, Lingulodinium machaerophorum strongly dominates the associations which are additionally characterized by cysts of Gymnodinium nolleri, cysts of Polykrikos kofoidii and cysts of Polykrikos schwartzii. (2) Off Cape Yubi, a region with increasing perennial upwelling, L. machaerophorum, Brigantedinium spp., species of the genus Impagidinium and cysts of Protoperidinium stellatum occur in highest relative abundances. (3) In coastal samples between Cape Ghir and Cape Yubi, Gymnodinium catenatum, species of the genus Impagidinium, Nematosphaeropsis labyrinthus, Operculodinium centrocarpum, cysts of P. stellatum and Selenopemphix nephroides determine the species composition. (4) Off Cape Blanc, where upwelling prevails perennially, and at offshore sites, heterotrophic dinocyst species show highest relative abundances. A Redundancy Analysis reveals fluvial mud, sea surface temperature and the depth of the mixed layer in boreal spring (spring) as the most important parameters relating to the dinocyst species association. Dinocyst accumulation rates were calculated for a subset of samples using well-constrained sedimentation rates. Highest accumulation rates with up to almost 80.000 cysts cm**-2 ky**-1 were found off Cape Ghir and Cape Yubi reflecting their eutrophic upwelling filaments. A Redundancy Analysis gives evidence that primary productivity and the input of fluvial mud are mostly related to the dinocyst association. By means of accumulation rate data, quantitative cyst production of individual species can be considered independently from the rest of the association, allowing autecological interpretations. We show that a combined interpretation of relative abundances and accumulation rates of dinocysts can lead to a better understanding of the productivity conditions off NW Africa.

Abundance of organic-walled dinoflagellate cysts of sediment core GeoB3603-2

Here, we reconstruct the varying influence of the Agulhas Current (AgC), the South Atlantic Current and the Antarctic Circumpolar Current on the Agulhas Retroflection (AgR) in the eastern South Atlantic Ocean for the last 160,000 years on the basis of the dinoflagellate cysts, pollen and spores present in a sediment core (GeoB 3603-2) from the southeastern Cape Basin offshore South Africa, where the Agulhas Current enters the Atlantic Ocean. Our analyses reveal strong orbital forcing on the heat exchange between the Indian Ocean and the South Atlantic Ocean during the Late Quaternary. Maxima in local productivity appear to be primarily related to a strengthening of the ocean circulation as a result of the high seasonal contrast during precession maxima. During precession minima, seasonal contrast was low and stratified, oligo- to mesotrophic conditions prevailed, notably when these minima coincided with the glacial terminations. The clear presence of periodicities on a sub-Milankovitch scale as well as modulations of the primary frequencies demonstrate that the Agulhas Retroflection furthermore is modulated substantially by complex interactions between the subtropical, 'precession-driven', climate and ocean circulation systems, and the southern, 'obliquity-driven', high latitudes.

Dinoflagellate cysts in western equatorial Atlantic surface sediments

In contrast to the wide range of studies carried out in temperate and high-latitude oceanic regions, only a few studies have focused on recent and Holocene organic-walled dinoflagellate cyst assemblages from the tropics. This information is, however, essential for fully understanding the ability of species to adapt to different oceanographic regimes, and ultimately their potential application to local and regional palaeoenvironmental and palaeoceanographic reconstructions. Surface sediment samples of the western equatorial Atlantic Ocean north of Brazil, an area greatly influenced by Amazon River discharge waters, were therefore analysed in detail for their organic-walled dinoflagellate cyst content. A diverse association of 43 taxa was identified, and large differences in cyst distribution were observed. The cyst thanatocoenosis in bottom sediments reflects the seasonal advection of Amazon River discharge water through the Guyana Current and the North Equatorial Countercurrent well into the North Atlantic. To establish potential links between cyst distribution and the environmental conditions of the upper water column, distribution patterns were compared with mean temperature, salinity, density and stratification gradients within the upper water column (0-100 m) over different times of the year, using correspondence analysis and canonical correspondence analysis. The analyses show that differences in these parameters only play a subordinate role in determining species distribution. Instead, nutrient availability, or related factors, dominates the distribution pattern. The only possible indicators of slightly reduced salinities are Trinovantedinium applanatum and Lingulodinium machaerophorum. Four assemblage groups of cyst taxa with similar environmental affinities related to specific water masses/currents can be distinguished and have potential for palaeoenvironmental reconstruction.

(Appendix A) Radiolarian abundance in ODP Hole 202-1241A

A quantitative radiolarian study at Ocean Drilling Program Site 1241 in the eastern tropical Pacific enables us to reconstruct paleoceanographic changes that occurred since the latest middle Miocene. Today, this site is located just under the Eastern Pacific Warm Pool (EPWP). Based on the abundance variations of radiolarian characteristic species which are indicators of upwelling and thermocline changes, it is suggested that three notable changes occurred at 10.6, 9.8, and 4.2 Ma in the region. Four distinct periods of oceanographic conditions bounded by these notable changes were characterized on the basis of the following: (1) stratified seawater (12.0 to 10.6 Ma); (2) a shallowing of the thermocline and an increasing of upwelling (10.6 to 9.8 Ma); (3) significant inflow of warm water to the eastern tropical Pacific caused by an intensified Northern Equatorial Countercurrent (NECC), resulting in the formation of EPWP (9.8 to 4.2 Ma); and (4) the reduction of the EPWP and the NECC, and an increase in upwelling (4.2 to 0 Ma). The timing of these paleoceanographic events indicated the strong relations with the opening and closing of the Indonesian and Central American (Panama) Seaways. The reduction of the EPWP (this study) and the deepening of the thermocline in western Pacific at about 4.2 Ma (Cannariato and Ravelo, 1997; Chaisson and Ravelo, 2000) indicated a change from a state resembling El Niño in the late Miocene and the early Pliocene time to a state resembling La Niña by the late Pliocene