Ornithogenic soils and the paleoecology of pygoscelid penguins in Antarctica

Ornithogenic or bird-formed soils have accumulated in many coastal regions around Antarctica as a result of breeding activities by pygoscelid penguins, especially the Adelie penguin (Pygoscelis adeliae). These soils are often deep, range from hundreds to thousands of years old, and contain a natural archive of penguin tissues and those of their prey. In some regions, these tissues are extremely well preserved by the dry, cold environment and include complete and partial penguin mummies, feathers, bone, and eggshell. Hard parts of prey (fish bones, otoliths, and squid beaks) also commonly occur in these deposits from the penguin guano as it accumulates during soil development. Here, we review how research on these soils and the tissues they contain has progressed since they were first identified and described. These studies have provided not only valuable information on penguin occupation history with climate change since the Pleistocene, but also whole ecosystem responses to perturbations such as the 'krill surplus' that is hypothesized to have occurred following historic depletion of seals and whales in the 18th-20th centuries. New findings in the Ross Sea indicate how penguin occupation and abandonment cycles have progressed over millennia in relation to climate change. In addition, stable isotope analysis of delta N-15 and delta C-13 in ancient and modern Adelie penguin tissues (feathers, bone, eggshell and membrane) and guano support the 'krill surplus' hypothesis in showing a dietary shift from fish to krill over the past similar to 200 years. Other recent studies have focused on stable isotope analyses of penguin prey remains, as well as ancient DNA and mercury analyses of penguin tissues recovered from ornithogenic soils. An analysis of fish otoliths recovered from ancient guano provide a means to investigate values of otolith carbonate delta O-18, which correlates with other paleoclimatic records, and can be used as a proxy for changing ocean temperatures through time. In addition, measurements of total mercury (Hg) in penguin egg membrane from abandoned colonies up to 800 years old indicate significantly higher mercury levels in the past compared to modern penguins, likely due to a greater reliance on higher trophic prey prior to the proposed 'krill surplus'. All of these studies indicate that ornithogenic soils and the natural archive of tissues they contain provide a unique means to integrate both terrestrial and marine records with ecosystem studies and climate change, past and present, in Antarctica. (C) 2014 Elsevier Ltd and INQUA. All rights reserved.