Please cite this article as: Pomar, L., Baceta, J.I., Hallock, P., Mateu-Vicens, G., Basso, D., Reef building and carbonate production modes in the west-central Tethys during the Cenozoic, Marine and Petroleum Geology (2017), ABSTRACTChanging components, rock textures, lithofacies, platform types and architecture throughout time are unique characteristics of carbonate rocks. Characterizing these attributes has been approached by 1) building reference models for specific Phanerozoic intervals, 2) recognizing the climatic impact in modulating carbonate production, and 3) analyzing the influence of changing bio-geochemical conditions. The reference-model approach is mostly based on biological evolution, the climaticimpact approach emphasizes temperature, and the bio-geochemical approach considers the changes in Mg/Ca ratios and Ca ++ concentrations in the oceans. To date, however, an analysis integrating all of these factors is still missing. The analysis presented here includes all these factors but also CO 2 , which is fundamental for both photosynthesis and CaCO 3 precipitation.Here we analyze the waxing and waning of Cenozoic reef limestones from the central Tethys region through several steps: 1) on the basis of rock volume, rock textures, associated sediments and light-dependent skeletal components, as records of light penetration and wave energy (depth); 2) on global environmental conditions (δ 13 C, δ 18 O, pCO 2 , temperature); and 3) on the basis of functionality, nutritional requirements and available resources.Through the Cenozoic, water motion, whether induced by surface or internal waves or by currents, increased as the thermal gradients strengthened, both with depth and with latitude. Active water motion is essential for plankton catchers such as corals, but less so for many larger benthic foraminifers (LBF). Pycnoclines in the meso-oligophotic zone would then favor the benthic plankton catchers such as corals, but would be detrimental for many LBF. Warm temperatures favored LBF. The Eocene LBF families predominated during lowering of atmospheric pCO 2 by using respiratory CO 2 to enhance the symbiont production of photosynthates under oligotrophic conditions and limited turbulence, whereas the Miocene families had to adapt to a progressive increase in turbulence. The eurythermal coralline red algae, however, became preponderant producers in the mesophotic zone during times when the δ 13 C was relatively high. This explains two apparent paradoxes: 1) corals thrive best when the Earth's high latitudes cool, and 2) the dominance of corals and LBF is inversely correlated, despite they both require tropical conditions and have similar trophic strategies (mixotrophy).

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The most important groups of modern red calcareous algae are the Mg-calcite secreting Corallinales and Sporolithales, and the aragonitic Peyssonneliales and Nemaliales. They are common on the world's shelves and are vulnerable to the global warming and the lowering of pH of sea water, caused by the ongoing increase in anthropogenic CO 2 . Among them, coralline algae are ecosystem engineers and major producers of carbonate sediment, of particular importance in temperate and cold seas. Corallines respond to marine acidification and rising temperature showing decreased net calcification, decreased growth and reproduction, as well as reduced abundance and diversity, leading to death and ecological shift to dominant non-calcifying algae. Despite their key ecological and sedimentological role, and because of their vulnerability to marine warming and acidification, our knowledge of the distribution of coralline-dominated habitats and the quantification of their carbonate production is not adequate to allow proper environmental management and confident modelling of a global carbon budget. Locating the algal carbonate factories around the world, then describing them, e.g., evaluating their extent and their production, are a priority for future research. résumé Production carbonatée par les algues rouges calcaires et changement climatique global.Les groupes les plus importants d'algues rouges calcaires actuelles sont, d'une part, les Corallinales et les Sporolithales, qui sécrètent de la calcite magnésienne, et d'autre part, les Peyssonnéliales et les Némaliales, qui produisent de l'aragonite. Abondantes sur les plates-formes actuelles, elles y jouent un rôle majeur

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ABSTRACT1. The protocols available for sampling and monitoring shallow subtidal rhodolith beds (RBs) are inadequate for the deep Mediterranean analogues, and need calibration in order to attain comparable results.2. After reviewing the present knowledge of the specificities of Mediterranean RBs, and in the framework of the ongoing international effort for their conservation, a two-step approach is suggested for their definition, identification, delimitation, description, and monitoring.3. Regional mapping should be improved, and RBs should be identified and delimited as those areas of the sea floor with >10% cover of live rhodoliths over a minimum surface of 500 m 2 , on 1:10000 scale. More detailed scales (at least 1:1000) should be used for monitoring selected RBs, in order to detect significant changes through time. 4. Beside location and areal extent, the description of RBs should include the occurrence of macroscopic sedimentary structures of the sea floor, thickness of live cover, mean percentage cover of live thalli and surface live/dead ratio, cover of dominant morphologies of rhodoliths (simplified on a ternary diagram), and volumetrically important calcareous algal species.5. For the purpose of assessment of the ecological status and the evaluation of human-induced impacts, quantitative data about community composition are required. The comparative assessment of ecological status and the identification of RBs of high conservation value for special protection should consider the natural geographic and seasonal/annual variability of RBs.

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Lithophyllum species in the Mediterranean Sea function as algal bioconstructors, contributing to the formation of biogenic habitats such as coralligenous concretions. In such habitats, thalli of Lithophyllum, consisting of crusts or lamellae with entire or lobed margins, have been variously referred to as either one species, L. stictiforme, or two species, L. stictiforme and L. cabiochiae, in the recent literature. We investigated species diversity and phylogenetic relationships in these algae by sequencing three markers (psbA and rbcL genes, cox2,3 spacer), in conjunction with methods for algorithmic delimitation of species (ABGD and GMYC). Mediterranean subtidal Lithophyllum belong to a well‐supported lineage, hereby called the L. stictiforme complex, which also includes two species described from the Atlantic, L. lobatum and L. searlesii. Our results indicate that the L. stictiforme complex consists of at least 13 species. Among the Mediterranean species, some are widely distributed and span most of the western and central Mediterranean, whereas others appear to be restricted to specific localities. These patterns are interpreted as possibly resulting from allopatric speciation events that took place during the Messinian Salinity Crisis and subsequent glacial periods. A partial rbcL sequence from the lectotype of L. stictiforme unambiguously indicates that this name applies to the most common subtidal Lithophyllum in the central Mediterranean. We agree with recent treatments that considered L. cabiochiae and L. stictiforme conspecific. The diversity of Lithophyllum in Mediterranean coralligenous habitats has been substantially underestimated, and future work on these and other Mediterranean corallines should use identifications based on DNA sequences.

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