High-resolution stratigraphic framework for Mediterranean sapropel S5

A high-resolution stratigraphic framework is presented for sapropel S5, which represents the low-mid latitude climate optimum of the previous interglacial period (Eemian). The framework is based on three sites along a transect from west to east through the eastern Mediterranean, and is further validated using a fourth site. This method allows expression of S5-based proxy records of Eemian climate variability along a standardised depth scale that offers unprecedented possibilities for assessment of spatial gradients and signal leads and lags in an interval where highresolution (radiocarbon-style) dating cannot be performed. Our lateral comparison of S5 sapropels suggests that the onset of S5 in ODP site 967C (Eratosthenes seamount) was 1-6 centuries delayed relative to the onsets in more westerly sites.

Species-specific consequences of ocean acidification for the calcareous tropical green algae Halimeda, 2011

Ocean acidification (OA), resulting from increasing dissolved carbon dioxide (CO2) in surface waters, is likely to affect many marine organisms, particularly those that calcify. Recent OA studies have demonstrated negative and/or differential effects of reduced pH on growth, development, calcification and physiology, but most of these have focused on taxa other than calcareous benthic macroalgae. Here we investigate the potential effects of OA on one of the most common coral reef macroalgal genera,Halimeda. Species of Halimeda produce a large proportion of the sand in the tropics and are a major contributor to framework development on reefs because of their rapid calcium carbonate production and high turnover rates. On Palmyra Atoll in the central Pacific, we conducted a manipulative bubbling experiment to investigate the potential effects of OA on growth, calcification and photophysiology of 2 species of Halimeda. Our results suggest that Halimeda is highly susceptible to reduced pH and aragonite saturation state but the magnitude of these effects is species specific. H. opuntiasuffered net dissolution and 15% reduction in photosynthetic capacity, while H. taenicola did not calcify but did not alter photophysiology in experimental treatments. The disparate responses of these species to elevated CO2 partial -pressure (pCO2) may be due to anatomical and physiological differences and could represent a shift in their relative dominance in the face of OA. The ability for a species to exert biological control over calcification and the species specific role of the carbonate skeleton may have important implications for the potential effects of OA on ecological function in the future.

Appendix 7: List of species with their distribution ranges restricted to the corresponding phytogeographical region

Understanding species distribution patterns and the corresponding environmental determinants is a crucial step in the development of effective strategies for the conservation and management of plant communities and ecosystems. Therefore, a central prerequisite is the biogeographical and macroecological analysis of factors and processes that determine contemporary, potential, as well as future geographic distribution of species. This thesis has been conducted in the framework of the BIOMAPS-BIOTA project at the Nees Institute of Biodiversity of Plants, which was funded by the German Federal Ministry of Education and Research (BMBF). The study investigated patterns of plants species richness and phytogeographic regions under contemporary environmental conditions and forecasted future climate change in the area of West Africa covering five countries: Benin, Burkina Faso, Côte d'Ivoire, Ghana and Togo. Firstly, geographic patterns of vascular plant species richness have been depicted at a relatively fine spatial resolution based on the potential distribution of 3,393 species. Species richness is closely related to the steep climatic gradient existing in the region with a high concentration of species in the most humid areas in the south and decreases towards the northern drier areas. The investigation of the effectiveness of the existing network of protected areas shows an overall good coverage of species in the study area. However, the proportion of covered species is considerably lower at national extent for some countries, thus calling for more protected areas in order to cover adequately a maximum number of plants species in these countries. Secondly, based on the potential distribution range of vascular plant species, seven phytogeographic regions have been delineated that broadly reflect the vegetation zones as defined by White (1983). However notable differences to the delineation of White (1983) occur at the margins of some regions. Corresponding to a general southward shifted of all regions. And expansion of the Sahel vegetation zone is observed in the north, while the rainforest zone is decreased in the very south.This is alarming since the rainforest shelters a high number of species and a high proportion of range-restricted or endemic species, despite their relatively small extent compared to the other regions. Finally, the evaluation of the potential impact of climate change on plant species richness in the study area, results in a severe loss of future suitable habitat for up to 50% of species per grid cell, particularly in the rainforest region. Moreover, the analysis of the possible shift of phytogeographic regions shows in general a strong deterioration of the West African rainforest. In contrast the drier areas are expanding continuously, although a slight gain in species number can be observed in some particular regions. The overall lesson to retain from the results of this study is that the West African rainforest should be fixed as a high priority area for the conservation of biodiversity of plants, since it is subject to severe contemporary and projected future threats.

Differential response of two Mediterranean cold-water coral species to ocean acidification

Cold-water coral (CWC) reefs constitute one of the most complex deep-sea habitats harboring a vast diversity of associated species. Like other tropical or temperate framework builders, these systems are facing an uncertain future due to several threats, such as global warming and ocean acidification. In the case of Mediterranean CWC communities, the effect may be exacerbated due to the greater capacity of these waters to absorb atmospheric CO2 compared to the global ocean. Calcification in these organisms is an energy-demanding process, and it is expected that energy requirements will be greater as seawater pH and the availability of carbonate ions decrease. Therefore, studies assessing the effect of a pH decrease in skeletal growth, and metabolic balance are critical to fully understand the potential responses of these organisms under a changing scenario. In this context, the present work aims to investigate the medium- to long-term effect of a low pH scenario on calcification and the biochemical composition of two CWCs from the Mediterranean, Dendrophyllia cornigera and Desmophyllum dianthus. After 314 d of exposure to acidified conditions, a significant decrease of 70 % was observed in Desmophyllum dianthus skeletal growth rate, while Dendrophyllia cornigera showed no differences between treatments. Instead, only subtle differences between treatments were observed in the organic matter amount, lipid content, skeletal microdensity, or porosity in both species, although due to the high variability of the results, these differences were not statistically significant. Our results also confirmed a heterogeneous effect of low pH on the skeletal growth rate of the organisms depending on their initial weight, suggesting that those specimens with high calcification rates may be the most susceptible to the negative effects of acidification.

Coccolithophore sensitivities to changing carbonate chemistry - an ecological framework

Coccolithophores are a group of unicellular phytoplankton species whose ability to calcify has a profound influence on biogeochemical element cycling. Calcification rates are controlled by a large variety of biotic and abiotic factors. Among these factors, carbonate chemistry has gained considerable attention during the last years as coccolithophores have been identified to be particularly sensitive to ocean acidification. Despite intense research in this area, a general concept harmonizing the numerous and sometimes (seemingly) contradictory responses of coccolithophores to changing carbonate chemistry is still lacking to date. Here, we present the "substrate-inhibitor concept" which describes the dependence of calcification rates on carbonate chemistry speciation. It is based on observations that calcification rate scales positively with bicarbonate (HCO3-), the primary substrate for calcification, and carbon dioxide (CO2), which can limit cell growth, whereas it is inhibited by protons (H+). This concept was implemented in a model equation, tested against experimental data, and then applied to understand and reconcile the diverging responses of coccolithophorid calcification rates to ocean acidification obtained in culture experiments. Furthermore, we (i) discuss how other important calcification-influencing factors (e.g. temperature and light) could be implemented in our concept and (ii) embed it in Hutchinson's niche theory, thereby providing a framework for how carbonate chemistry-induced changes in calcification rates could be linked with changing coccolithophore abundance in the oceans. Our results suggest that the projected increase of H+ in the near future (next couple of thousand years), paralleled by only a minor increase of inorganic carbon substrate, could impede calcification rates if coccolithophores are unable to fully adapt. However, if calcium carbonate (CaCO3) sediment dissolution and terrestrial weathering begin to increase the oceans' HCO3- and decrease its H+ concentrations in the far future (10 -100 kyears), coccolithophores could find themselves in carbonate chemistry conditions which may be more favorable for calcification than they were before the Anthropocene.

Sponge bioerosion accelerated by ocean acidification across species and latitudes?

In many marine biogeographic realms, bioeroding sponges dominate the internal bioerosion of calcareous substrates such as mollusc beds and coral reef framework. They biochemically dissolve part of the carbonate and liberate so-called sponge chips, a process that is expected to be facilitated and accelerated in a more acidic environment inherent to the present global change. The bioerosion capacity of the demosponge Cliona celata Grant, 1826 in subfossil oyster shells was assessed via alkalinity anomaly technique based on 4 days of experimental exposure to three different levels of carbon dioxide partial pressure (pCO2) at ambient temperature in the cold-temperate waters of Helgoland Island, North Sea. The rate of chemical bioerosion at present-day pCO2 was quantified with 0.08-0.1 kg/m**2/year. Chemical bioerosion was positively correlated with increasing pCO2, with rates more than doubling at carbon dioxide levels predicted for the end of the twenty-first century, clearly confirming that C. celata bioerosion can be expected to be enhanced with progressing ocean acidification (OA). Together with previously published experimental evidence, the present results suggest that OA accelerates sponge bioerosion (1) across latitudes and biogeographic areas, (2) independent of sponge growth form, and (3) for species with or without photosymbionts alike. A general increase in sponge bioerosion with advancing OA can be expected to have a significant impact on global carbonate (re)cycling and may result in widespread negative effects, e.g. on the stability of wild and farmed shellfish populations, as well as calcareous framework builders in tropical and cold-water coral reef ecosystems.

Hidden impacts of ocean acidification to live and dead coral framework

Cold-water corals, such as Lophelia pertusa, are key habitat-forming organisms found throughout the world's oceans to 3000 m deep. The complex three-dimensional framework made by these vulnerable marine ecosystems support high biodiversity and commercially important species. Given their importance, a key question is how both the living and the dead framework will fare under projected climate change. Here, we demonstrate that over 12 months L. pertusa can physiologically acclimate to increased CO2, showing sustained net calcification. However, their new skeletal structure changes and exhibits decreased crystallographic and molecular-scale bonding organization. Although physiological acclimatization was evident, we also demonstrate that there is a negative correlation between increasing CO2 levels and breaking strength of exposed framework (approx. 20-30% weaker after 12 months), meaning the exposed bases of reefs will be less effective 'load-bearers', and will become more susceptible to bioerosion and mechanical damage by 2100.

Morphometric measurements throughout ontogeny of selected planktic foraminiferal test species

Development plays an important part in shaping adult morphology and morphological disparity, yet its influence on evolutionary processes is seldom explored because of a lack of preservation of ontogenetic stages in the fossil record. By preserving their entire ontogenetic history within their test, and with the advent of high-resolution imaging techniques, planktic foraminifera allow us to investigate the influence of developmental constraints on disparity. Using Synchrotron radiation X-ray tomographic microscopy (SRXTM), we reconstruct the ontogenetic progression of seven species across several of the major morphotypic groups of planktic foraminifera, including morphotypes of a species exhibiting high phenotypic plasticity and closely related pseudo-cryptic sister-taxa. We show differences in growth patterns between the globigerinid species, which appear more tightly regulated within the framework of isometry from the neanic stage, and the globorotaliid species, whose adult stages present allometric trends. Morphological change through ontogeny results in a change in surface area to volume ratios. Different metabolic processes therefore dominate at different stages of ontogeny, changing the vulnerability of the organism to environmental influences over growth, from factors affecting diffusion rates in the juvenile to those affecting energy supply in the adult. These findings identify some of the parameters within which evolutionary mechanisms have to act.

Georeferenced standardized apparent wave power along the coastline of Cabrera National Park for analysis of essential habitats of littoral species

Wind- induced exposure is one of the major forces shaping the geomorphology and biota in coastal areas. The effect of wave exposure on littoral biota is well known in marine environments (Ekebon et al., 2003; Burrows et al., 2008). In the Cabrera Archipelago National Park wave exposure has demostrated to have an effect on the spatial distribution of different stages of E.marginatus (Alvarez et al., 2010). Standarized average wave exposures during 2008 along the Cabrera Archipelago National park coast line were calculated to be applied in studies of littoral species distribution within the archipelago. Average wave exposure (or apparent wave power) was calculated for points located 50 m equidistant on the coastline following the EXA methodology (EXposure estimates for fragmented Archipelagos) (Ekebon et al., 2003). The average wave exposures were standardized from 1 to 100 (minimum and maximum in the area), showing coastal areas with different levels of mea wave exposure during the year. Input wind data (direction and intensity) from 2008 was registered at the Cabrera mooring located north of Cabrera Archipelago. Data were provided by IMEDEA (CSIC-UIB, TMMOS http://www.imedea.uib-csic.es/tmoos/boyas/). This cartography has been developed under the framework of the project EPIMHAR, funded by the National Park's Network (Spanish Ministry of Environment, Maritime and Rural Affairs, reference: 012/2007 ). Part of this work has been developed under the research programs funded by "Fons de Garantia Agrària i Pesquera de les Illes Balears (FOGAIBA)".