Bryoliths constructed by bryozoans in symbiotic associations with hermit crabs, Golfe d'Arguin, Mauritania

The Golfe d'Arguin offshore of northern Mauritania hosts a rare modern analogue for heterozoan carbonate production in a tropical marine setting. Dominated by ocean upwelling and with additional fertilisation by iron-rich aeolian dust, this naturally eutrophic marine environment lacks typical photozoan communities. A highly productive, tropical cosmopolitan biota dominated by molluscs and suspension-feeders such as bryozoans and balanids characterises the carbonate-rich surface sediments. Overall biodiversity is relatively low and the species present are tolerant against the eutrophic and low-light conditions, the strong hydrodynamic regime governed by ocean upwelling, and the unstable, soft-bottom seafloor with few hard substrata. Here, we describe an ectosymbiosis between the hermit crab Pseudopagurus granulimanus (Miers, 1881) and monospecific assemblages of the encrusting cheilostome bryozoan Acanthodesia commensale (Kirkpatrick and Metzelaar, 1922) that cohabits vacant gastropod shells. Nucleating on an empty gastropod shell, the bryozoan colonies form multilamellar skeletal crusts that produce spherical encrustations and extend the living chamber of the hermit crab through helicospiral tubular growth. This non-obligate mutualistic symbiosis illustrates the adaptive capabilities and benefits from a close partnership in a complex marine environment, driven by trophic conditions, high water energies and instable substratum. Sectioned bryoliths show that between 49 and 97 % of the solid volume of the specimens consists of bryozoan skeleton.

Globale räumlich-zeitliche Klimavariabilität im Holozän (GHOST) [Global Holocene Spatial and Temporal Climate Variability]

Verbindung mariner Paläotemperatur-Kurven mit dreidimensionaler, gekoppelter Atmosphäre-Ozean Modellierung [Integrating marine multiproxy temperature estimates and three-dimensional coupled atmosphere/ocean modelling] Das Projekt war ein Beitrag zur Untersuchung des Klimas des Holozäns. Es basierte auf zwei Standbeinen: Der Heranziehung von weltweit verfügbaren, unbearbeiteten, aktualisierten und neu zusammengestellten marinen multiproxy Temperaturrekonstruktionen einerseits und der Verwendung von gekoppelten Zirkulationsmodellen für Atmosphäre und Ozean andererseits. Das Modell arbeitete mit relativ geringer Auflösung und Rechenzeit und ist für transiente Simulationen des Paläoklimas angepaßt. Für eine möglichst große globale Abdeckung der Zeitserien von Klimaproxies wurden Sedimentdaten herangezogen, die eine geringe aber dennoch höchstmögliche zeitliche Auflösung im Bereich von 50 bis 200 Jahren besitzen. Sowohl Datenrekonstruktion als auch gekoppelte Klimamodellierung erzeugten dreidimensionale Datensätze, zwei räumliche Dimensionen auf der Erdoberfläche, sowie die Zeit als dritte Dimension. Raumzeitliche Muster wurden im Rahmen des Projektes untersucht. Die eingehende Analyse rekonstruierter wie der Modell-Daten sollte einerseits das Verständnis für Klimaänderungen verbessern, die in Proxydaten gefunden werden und andererseits eine Validierung der Klimavariabilität im Modell ermöglichen. Die Musteranalyse ergab Einblicke in die Mechanismen, die zur Heterogenität von Erwärmung und Abkühlung im Holozän beitragen. Die Weiterführung der Klimasimulationen des Holozäns in die Zukunft der nächsten Jahrhunderte diente einer besseren Abschätzung der zukünftigen Klimaänderung.

Seawater carbonate chemistry and processes during experiments with crabs Chionoecetes tanneri and Cancer magister, 2007

Rising levels of atmospheric carbon dioxide could be curbed by large-scale sequestration of CO2 in the deep sea. Such a solution requires prior assessment of the impact of hypercapnic, acidic seawater on deep-sea fauna. Laboratory studies were conducted to assess the short-term hypercapnic tolerance of the deep-sea Tanner crab Chionoecetes tanneri, collected from 1000 m depth in Monterey Canyon off the coast of central California, USA. Hemolymph acid- base parameters were monitored over 24 h of exposure to seawater equilibrated with ~1% CO2 (seawater PCO2 ~6 torr or 0.8 kPa, pH 7.1), and compared with those of the shallow-living Dungeness crab Cancer magister. Short-term hypercapnia-induced acidosis in the hemolymph of Chionoecetes tanneri was almost uncompensated, with a net 24 h pH reduction of 0.32 units and a net bicarbonate accumulation of only 3 mM. Under simultaneous hypercapnia and hypoxia, short-term extracellular acidosis in Chionoecetes tanneri was completely uncompensated. In contrast, Cancer magister fully recovered its hemolymph pH over 24 h of hypercapnic exposure by net accumulation of 12 mM bicarbonate from the surrounding medium. The data support the hypothesis that deep-sea animals, which are adapted to a stable environment and exhibit reduced metabolic rates, lack the short-term acid-base regulatory capacity to cope with the acute hypercapnic stress that would accompany large-scale CO2 sequestration. Additionally, the data indicate that sequestration in oxygen-poor areas of the ocean would be even more detrimental to deep-sea fauna.

CO2-driven decrease in pH disrupts olfactory behaviour and increases individual variation in deep-sea hermit crabs

Deep-sea species are generally thought to be less tolerant of environmental variation than shallow-living species due to the relatively stable conditions in deep waters for most parameters (e.g. temperature, salinity, oxygen, and pH). To explore the potential for deep-sea hermit crabs (Pagurus tanneri) to acclimate to future ocean acidification, we compared their olfactory and metabolic performance under ambient (pH 7.6) and expected future (pH 7.1) conditions. After exposure to reduced pH waters, metabolic rates of hermit crabs increased transiently and olfactory behaviour was impaired, including antennular flicking and prey detection. Crabs exposed to low pH treatments exhibited higher individual variation for both the speed of antennular flicking and speed of prey detection, than observed in the control pH treatment, suggesting that phenotypic diversity could promote adaptation to future ocean acidification.

Expanded GHOST database

In this study we review a global set of alkenone- and foraminiferal Mg/Ca-derived sea surface temperatures (SST) records from the Holocene and compare them with a suite of published Eemian SST records based on the same approach. For the Holocene, the alkenone SST records belong to the actualized GHOST database (Kim, J.-H., Schneider R.R., 2004). The actualized GHOST database not only confirms the SST changes previously described but also documents the Holocene temperature evolution in new oceanic regions such as the Northwestern Atlantic, the eastern equatorial Pacific, and the Southern Ocean. A comparison of Holocene SST records stemming from the two commonly applied paleothermometry methods reveals contrasting - sometimes divergent - SST evolution, particularly at low latitudes where SST records are abundant enough to infer systematic discrepancies at a regional scale. Opposite SST trends at particular locations could be explained by out-of-phase trends in seasonal insolation during the Holocene. This hypothesis assumes that a strong contrast in the ecological responses of coccolithophores and planktonic foraminifera to winter and summer oceanographic conditions is the ultimate reason for seasonal differences in the origin of the temperature signal provided by these organisms. As a simple test for this hypothesis, Eemian SST records are considered because the Holocene and Eemian time periods experienced comparable changes in orbital configurations, but had a higher magnitude in insolation variance during the Eemian. For several regions, SST changes during both interglacials were of a similar sign, but with higher magnitudes during the Eemian as compared to the Holocene. This observation suggests that the ecological mechanism shaping SST trends during the Holocene was comparable during the penultimate interglacial period. Although this "ecology hypothesis" fails to explain all of the available results, we argue that any other mechanism would fail to satisfactorily explain the observed SST discrepancies among proxies.

Temperature and salinity reconstruction for sediment core GeoB3313-1

We reconstructed changes of temperature, salinity, and productivity within the southern Peru-Chile Current during the last 8000 years from a high-resolution sediment core recovered at 41°S using alkenones, isotope ratios of planktic foraminifera, biogenic opal, and organic carbon. Paleotemperatures and paleosalinities reached maximum values at ~5500 years ago and thereafter declined to modern values, whereas paleoproductivity continuously increased throughout the last 8000 years. We ascribe these long-term Holocene trends primarily to latitudinal shifts of the Antarctic Circumpolar Current (ACC). The concurrence with shifts in the position of the Southern Westerlies points to a common response of atmospheric and oceanographic circulation patterns off southern Chile. Millennial- to centennial-scale fluctuations of paleotemperatures and paleosalinities, on the other hand, lag displacements in the position of the Southern Westerlies but reveal a significant correlation to short-term temperature changes in Antarctica, indicating a high-latitude control of the ACC at these timescales.

Sea-surface tempertaure reconstruction of sediment cores from the Southeast Pacific

Applying the alkenone method, we estimated sea-surface temperatures (SSTs) for the past 33 kyr in two marine sediment cores recovered from the continental slope off mid-latitude Chile. The SST record shows an increase of 6.7°C from the last ice age (LIA) to the Holocene climatic optimum, while the temperature contrast between LIA and modern temperatures is only about 3.4°C. The timing and magnitude of the last deglacial warming in the ocean correspond to those observed in South American continental records. According to our SST record, the existence of a Younger Dryas equivalent cooling in the Southeast Pacific is much more uncertain than for the continental climate changes. A warming step of about 2.5°C observed between 8 and 7.5 cal kyr BP may have been linked to the early to mid-Holocene climatic transition (8.2-7.8 cal kyr BP), also described from equatorial Africa and Antarctica. In principal, variations in the latitudinal position of the Southern Pacific Westerlies are considered to be responsible for SST changes in the Peru-Chile current off mid-latitude Chile.

Haemolymph inorganic ion composition, physiology and behaviour dependent on temperature and ambient magnesium concentration in early life history stages of Paralomis granulosa (Decapoda, Anomura, Lithodidae)

Marine brachyuran and anomuran crustaceans are completely absent from the extremely cold (-1.8 °C) Antarctic continental shelf, but caridean shrimps are abundant. This has at least partly been attributed to low capacities for magnesium excretion in brachyuran and anomuran lithodid crabs ([Mg2+]HL = 20-50 mmol/L) compared to caridean shrimp species ([Mg2+]HL = 5-12 mmol/L). Magnesium has an anaesthetizing effect and reduces cold tolerance and activity of adult brachyuran crabs. We investigated whether the capacity for magnesium regulation is a factor that influences temperature-dependent activity of early ontogenetic stages of the Sub-Antarctic lithodid crab Paralomis granulosa. Ion composition (Na+, Mg2+, Ca2+, Cl-, [SO4]2-) was measured in haemolymph withdrawn from larval stages, the first and second juvenile instars (crabs I and II) and adult males and females. Magnesium excretion improved during ontogeny, but haemolymph sulphate concentration was lowest in the zoeal stages. Neither haemolymph magnesium concentrations nor Ca2+:Mg2+ ratios paralleled activity levels of the life stages. Long-term (3 week) cold exposure of crab I to 1 °C caused a significant rise of haemolymph sulphate concentration and a decrease in magnesium and calcium concentrations compared to control temperature (9 °C). Spontaneous swimming activity of the zoeal stages was determined at 1, 4 and 9 °C in natural sea water (NSW, [Mg2+] = 51 mmol/L) and in sea water enriched with magnesium (NSW + Mg2+, [Mg2+] = 97 mmol/L). It declined significantly with temperature but only insignificantly with increased magnesium concentration. Spontaneous velocities were low, reflecting the demersal life style of the zoeae. Heart rate, scaphognathite beat rate and forced swimming activity (maxilliped beat rate, zoea I) or antennule beat rate (crab I) were investigated in response to acute temperature change (9, 6, 3, 1, -1 °C) in NSW or NSW + Mg2+. High magnesium concentration reduced heart rates in both stages. The temperature-frequency curve of the maxilliped beat (maximum: 9.6 beats/s at 6.6 °C in NSW) of zoea I was depressed and shifted towards warmer temperatures by 2 °C in NSW + Mg2+, but antennule beat rate of crab I was not affected. Magnesium may therefore influence cold tolerance of highly active larvae, but it remains questionable whether the slow-moving lithodid crabs with demersal larvae would benefit from an enhanced magnesium excretion in nature.

Haemolymph inorganic ion composition, physiology and behaviour dependent on temperature and magnesium in adult Paralomis granulosa (Decapoda, Anomura, Lithodidae)

A low capacity for regulation of extracellular Mg2+ has been proposed to exclude reptant marine decapod crustaceans from temperatures below 0°C and thus to exclude them from the high Antarctic. To test this hypothesis and to elaborate the underlying mechanisms in the most cold-tolerant reptant decapod family of the sub-Antarctic, the Lithodidae, thermal tolerance was determined in the crab Paralomis granulosa (Decapoda, Anomura, Lithodidae) using an acute stepwise temperature protocol (-1°, 1°, 4°, 7°, 10°, and 13°C). Arterial and venous oxygen partial pressures (Po2) in hemolymph, heartbeat and ventilation beat frequencies, and hemolymph cation composition were measured at rest and after a forced activity (righting) trial. Scopes for heartbeat and ventilation beat frequencies and intermittent heartbeat and scaphognathite beat rates at rest were evaluated. Hemolymph [Mg2+] was experimentally reduced from 30 mmol/L to a level naturally observed in Antarctic caridean shrimps (12 mmol/L) to investigate whether the animals remain more active and tolerant to cold (-1°, 1°, and 4°C). In natural seawater, righting speed was significantly slower at -1° and 13°C, compared with acclimation temperature (4°C). Arterial and venous hemolymph Po2 increased in response to cooling even though heartbeat and ventilation beat frequencies as well as scopes decreased. At rest, ionic composition of the hemolymph was not affected by temperature. Activity induced a significant increase in hemolymph [K+] at -1° and 1°C. Reduction of hemolymph [Mg2+] did not result in an increase in activity, an increase in heartbeat and ventilation beat frequencies, or a shift in thermal tolerance to lower temperatures. In conclusion, oxygen delivery in this cold-water crustacean was not acutely limiting cold tolerance, and animals may have been constrained more by their functional capacity and motility. In contrast to earlier findings in temperate and subpolar brachyuran crabs, these constraints remained insensitive to changing Mg2+ levels.

Epibiosis between the sand crab Emerita analoga and the mussel Semimytilus algosus

Coastal zones of the Humboldt Current Upwelling System (HCUS) are composed both of rocky and sandy beaches inhabited by macrozoobenthic communities. These show oscillating changes in the dominance of species; the abundance of the sand crab Emerita analoga is linked to phases of the El Niño Southern Oscillation (ENSO). The biogenic surfaces of these crabs serve as substrate for opportunistic colonizers. This study is the first record of an epibiosis between E. analoga and the rock mussel Semimytilus algosus, detected at a southern Peruvian sandy beach. Mussels fouled a wide size-range of adult E. analoga (7.3%) but they themselves belonged to small-size classes. The largest S. algosus was 17.4 mm in length. Highest permanence of epibionts was found on larger sand crabs (maximum between 24 and 27 mm). Significantly more mussels were found on the ventral surface (39.4%) compared to 10 other surface areas of the sand crab. Possible benefits and disadvantages of the observed epibiosis for both the basibiont and the epibiont are discussed.

Growth rates of late Miocene corals from Crete (Greece)

Modern scleractinian corals are classical components of marine shallow warm water ecosystems. Their occurrence and diversity patterns in the geological record have been widely used to infer past climates and environmental conditions. Coral skeletal composition data reflecting the nature of the coral environment are often affected by diagenetic alteration. Ghost structures of annual growth rhythms are, however, often well preserved in the transformed skeleton. We show that these relicts represent a valuable source of information on growth conditions of fossil corals. Annual growth bands were measured in massive hemispherical Porites of late Miocene age from the island of Crete (Greece) that were found in patch reefs and level bottom associations of attached mixed clastic environments as well as isolated carbonate environments. The Miocene corals grew slowly, about 2-4 mm/yr, compatible with present-day Porites from high-latitude reefs. Slow annual growth of the Miocene corals is in good agreement with the position of Crete at the margin of the Miocene reef belt. Within a given time slice, extension rates were lowest in level bottom environments and highest in attached inshore reef systems. Because sea surface temperatures (SSTs) can be expected to be uniform within a time slice, spatial variations in extension rates must reflect local variations in light levels (low in the level bottom communities) and nutrients (high in the attached reef systems). During the late Miocene (Tortonian-early Messinian), maximum linear extension rates remained remarkably constant within seven chronostratigraphic units, and if the relationship of SSTs and annual growth rates observed for modern massive Indo-Pacific Porites spp. applies to the Neogene, minimum (winter) SSTs were 20°-21°C. Although our paleoclimatic record has a low resolution, it fits the trends revealed by global data sets. In the near future we expect this new and easy to use Porites thermometer to add important new information to our understanding of Neogene climate.