The Diet of Early Birds Based on Modern and Fossil Evidence and a New
Framework for its Reconstruction
Abstract
Birds are some of the most diverse organisms on Earth, with species
inhabiting nearly every conceivable niche in every major biome. As such,
birds are vital to our understanding of modern ecosystems.
Unfortunately, this is hampered by knowledge gaps relating to the origin
of this modern diversity and its role in ecosystems. A crucial part of
addressing these shortcomings is improving our understanding of the
earliest birds, the non-avian avialans i.e. non-crown birds. The diet of
non-avian avialans has been a matter of substantial debate, partly
related to some of the ambiguous qualitative approaches that have been
used to reconstruct it. Here we review the methods of determining diet
in both modern avians and fossil avian and non-avian theropods, and
comment on their usefulness when applied to non-avian avialans. We use
this to propose a set of comparable, quantitative approaches to
ascertain fossil bird diet and on this basis provide a consensus of what
we currently know about fossil bird diet. While no single approach can
precisely predict diet in birds, each can exclude some diets and narrow
the dietary possibilities. We recommend combining [1] dental
microwear, [2] landmark-based muscular reconstruction, [3]
stable isotope geochemistry, [4] body mass estimations, [5]
traditional and/or geometric morphometric analysis, and [6] finite
element analysis to accurately reconstruct fossil bird diet. Our review
provides specific methodologies to implement each approach and discusses
complications future researchers should keep in mind. On this basis we
report the current state of knowledge of non-avian avialan diet which
remains very incomplete. The ancestral dietary condition in non-avian
avialans remains unclear due to a scarcity of data and contradictory
evidence in Archaeopteryx. Among early non-avian pygostylians,
Confuciusornis has finite element analysis and mechanical advantage
evidence pointing to herbivory, whilst Sapeornis only has mechanical
advantage evidence indicating granivory, which agrees with fossilised
ingested material known for this taxon. The enantiornithine
ornithothoracine Shenqiornis has mechanical advantage and pedal
morphometric evidence pointing to carnivory. In the hongshanornithid
ornithuromorph Hongshanornis, only mechanical advantage evidence
indicates granivory, but this is congruent with evidence of fossilised
ingested material in this taxon. The same is true for the
songlingornithid ornithuromorph Yanornis and its inferred carnivorous
diet. Due to the sparsity of robust dietary assignments, no clear trends
in non-avian avialan dietary evolution have yet emerged. Dietary
diversity may seem to increase through time, but this is a
preservational bias associated with a predominance of data from the
Early Cretaceous Jehol Lagerstatte. With this new framework and our
current synthesis of current knowledge of non-avian non-avialan diet, we
expect dietary knowledge and evolutionary trends to become much
clearer[…]