Atmosphere, ecology and evolution: what drove the Miocene expansion of C4 grasslands?

Grasses using the C4 photosynthetic pathway dominate today's savanna ecosystems and account for ∼20% of terrestrial carbon fixation. However, this dominant status was reached only recently, during a period of C4 grassland expansion in the Late Miocene and Early Pliocene (4–8 Myr ago). Declining atmospheric CO2 has long been considered the key driver of this event, but new geological evidence casts doubt on the idea, forcing a reconsideration of the environmental cues for C4 plant success.Here, I evaluate the current hypotheses and debate in this field, beginning with a discussion of the role of CO2 in the evolutionary origins, rather than expansion, of C4 grasses. Atmospheric CO2 starvation is a plausible selection agent for the C4 pathway, but a time gap of around 10 Myr remains between major decreases in CO2 during the Oligocene, and the earliest current evidence of C4 plants.An emerging ecological perspective explains the Miocene expansion of C4 grasslands via changes in climatic seasonality and the occurrence of fire. However, the climatic drivers of this event are debated and may vary among geographical regions.Uncertainty in these areas could be reduced significantly by new directions in ecological research, especially the discovery that grass species richness along rainfall gradients shows contrasting patterns in different C4 clades. By re-evaluating a published data set, I show that increasing seasonality of rainfall is linked to changes in the relative abundance of the major C4 grass clades Paniceae and Andropogoneae. I propose that the explicit inclusion of these ecological patterns would significantly strengthen climate change hypotheses of Miocene C4 grassland expansion. Critically, they allow a new series of testable predictions to be made about the fossil record.Synthesis. This paper offers a novel framework for integrating modern ecological patterns into theories about the geological history of C4 plants

Palaeoenvironment of Eocene prodelta in Spitsbergen recorded by the trace fossil Phycosiphon incertum

Ichnological, sedimentological and geochemical analyses were conducted on the Eocene Frysjaodden Formation in order to interpret palaeoenvironment prodelta sediments in the Central Basin of Spitsbergen. Phycosiphon incertum is the exclusive ichnotaxon showing differences in size, distribution, abundance and density, and relation to laminated/bioturbated intervals. Large P. incertum mainly occur dispersed, isolated and randomly distributed throughout the weakly laminated/non-laminated intervals. Small P. incertum occur occasionally in patches of several burrows within laminated intervals or as densely packed burrows in thin horizons in laminated intervals or constituting fully bioturbated intervals that are several centimetres thick. Ichnological changes are mainly controlled by oxygenation, although the availability of benthic food cannot be discarded. Changes in oxygenation and rate of sedimentation can be correlated with the registered variations in the Bouma sequence of the distal turbiditic beds within prodeltal shelf sediments.Funding for this research was provided by Project CGL2012-33281 (Secretaría de Estado de Investigación, Desarrollo e Innovación, Spain), Project RYC-2009-04316 (Ramón y Cajal Programme) and Projects RNM-3715 and RNM-7408 and Research Group RNM-178 (Junta de Andalucía). The authors benefited from a bilateral agreement between the universities of Granada and Oslo, supported by the University of Granada