One of the central measures used to describe modern ecosystems is the ratio of foliage area to that of the ground surface – the leaf area index (LAI) – which expresses the openness of vegetation canopies. A high LAI helps to retain moisture in the soil, partly by shading and cooling the surface to reduce evaporation and partly by stopping surface soil from being battered to a concrete-like consistency by heavy rain, which reduces the amount of water that can infiltrate. It is possible to estimate LAI across today’s entire land area using satellite image data but a proxy for palaeoecological LAI has remained hard to find.
The outer coating of leaves in well-shaded (high LAI) areas tends to have protective or pavement cells that are larger and have more complicated shapes than does that of leaves in more open canopies. The framework of leaf cells is silica-based and made up of structures known as phytoliths whose morphologies vary in much the same way as the cells that they support. So theoretically it is possible to use fossil phytoliths in terrestrial sediments to estimate LAI variations through time in local canopies, but first the approach needs a means of calibration from living ecosystems. The vegetation of Central American Costa Rica varies through the entire range of possible LAI values, which leads to varying amounts of sunlight available to the leaves of cover plants. Measuring the area and the degree of shape-complexity of phytoliths in modern soils there shows that each is positively correlated with LAI.
Putting this approach to use in the Cenozoic terrestrial sediments of Patagonia, US and Argentinean palaeoecologists aimed to examine how the evolution of the teeth of herbivorous mammals – a major feature in their speciation – linked to changes in vegetation structure (Dunn, R.E. et al. 2015. Linked canopy, climate and faunal change in the Cenozoic of Patagonia. Science, v. 347, p. 258-261). Using phytoliths they were able to show that in the Eocene the area was covered by dense, closed forest canopies that gradually became more open towards the end of the Eocene to be replaced by open forest and shrubland habitats in the Oligocene and Miocene, with a brief period of regreening. It was during the period of more open vegetation that tooth structure underwent the most change. Chances are that the vegetation shifts began in response to the onset of Antarctic glaciation at the beginning of the Oligocene Epoch and related climate change at the northern margin of the Southern Ocean. Changes in the herbivore teeth may have been in response to the increasing amount of dust adhering to leaves as canopies became more open and soil increasingly dried out.