Direct evidence of frugivory in the Mesozoic bird Longipteryx contradicts morphological proxies for diet

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Diet is one of the most important aspects of an animal's ecology, as it reflects direct interactions with other organisms and shapes morphology, behavior, and other life history traits. Modern birds (Neornithes) have a highly efficient and phenotypically plastic digestive system, allowing them to utilize diverse trophic resources, and digestive function has been put forth as a factor in the selectivity of the end-Cretaceous mass extinction, in which only neornithine dinosaurs survived.1 Although diet is directly documented in several early-diverging avian lineages,2 only a single specimen preserves evidence of diet in Enantiornithes, the dominant group of terrestrial Cretaceous birds.3 Morphology-based predictions suggest enantiornithines were faunivores,4,5,6 although the absence of evidence contrasts with the high preservation potential and relatively longer gut-retention times of these diets. Longipteryx is an unusual Early Cretaceous enantiornithine with an elongate rostrum; distally restricted dentition7; large, recurved, and crenulated teeth8; and tooth enamel much thicker than other paravians.9 Statistical analysis of rostral length, body size, and tooth morphology predicts Longipteryx was primarily insectivorous.4,5 Contrasting with these results, two new specimens of Longipteryx preserve gymnosperm seeds within the abdominal cavity interpreted as ingesta. Like Jeholornis, their unmacerated preservation and the absence of gastroliths indicate frugivory.10 As in Neornithes,11 complex diets driven by the elevated energetic demands imposed by flight, secondary rostral functions, and phylogenetic influence impede the use of morphological proxies to predict diet in early-diverging avian lineages.