Quantitative analysis of morphometric data of pre-modern birds: phylogenetic versus ecological signal

Birds are one of the most diverse clades of extant terrestrial vertebrates, a diversity that first arose during the Mesozoic as a multitude of lineages of pre-neornithine (stem) birds appeared but did not survive into the Cenozoic Era. Modern birds (Neornithes) inhabit an extensive array of ecologically distinct habitats and have specific and varied foraging strategies. Likewise, the morphological disparity among Mesozoic lineages appears to underscore a significant degree of ecological diversity, yet attempts to determine lineage specific ecologies have mainly been limited to superficial narratives. In recent years, numerous studies have used various morphometric proxies to interpret the paleoecology of Mesozoic bird lineages, but largely without evaluating the interplay between ecological and phylogenetic signals. Moreover, most studies of this sort transform the original data into logarithms to control dimensionality, underestimating the biases induced upon such transformations. The goal of this study is to quantitatively address the ecomorphology of crown-group Neornithes using a dense sample of raw forelimb and hindlimb measurements, and to examine if such results can be used to infer the ecologies of Mesozoic bird lineages. To that end, scaling of limb measurements and ecological data from modern birds was assessed statistically using phylogenetic comparative methods, followed by the inclusion of fossil taxa. A strong relationship was recovered between humerus and hindlimb allometric scaling and phylogeny. Our results indicate that while some ecological classes of modern birds can be discriminated from each other, phylogenetic signature can overwhelm ecological signal in morphometric data, potentially limiting the inferences that can be made from ecomorphological studies. Furthermore, we found differential scaling of leg bones among Early Cretaceous enantiornithines and ornithuromorphs, a result hinting that habitat partitioning among different lineages could be a pervasive phenomenon in avian evolutio

A theropod trackway providing evidence of a pathological foot from the exceptional locality of Las Hoyas (upper Barremian, Serrania de Cuenca, Spain)

We describe a trackway (LH-Mg-10-16) occurring in laminated carbonated limestones of the Las Hoyas locality, Serranı´a de Cuenca, Spain. It is unmistakably a large theropod dinosaur trackway encompassing two unusual aspects, namely, wide-steps, and a set of equally deformed left footprints (with a dislocated digit). The layer also preserves other vertebrate trails (fish Undichna) and different impressions in the sediment. To address these complex settings, we devised a multidisciplinary approach, including the ichnological and taphonomical descriptions, characterisation of the rock lithofacies using thin-sections, 3D structuredlight digitalisation with a high precision of 200–400 μm, and a geometric morphometric comparison with a large sample of bipedal dinosaur trackways. Sedimentary analyses showed that the trackway was produced in a humid, benthonic microbial mat, the consistency and plasticity of which enabled the preservation of the details of the movement of the animal. The results of the geometric analysis indicate that the “wide-steps” of the trackway is not unusual compared to other trackways, providing evidence that it was made by a single individual with an estimated hip height approximately 2 m. Analogous pathologies in extant archosaurs that yield the combination of wide steps and deformed digits in the same trackway were considered. All results mutually support the hypothesis that a large theropod dinosaur, with a pathological foot, generated the trackway as it crossed an area of shallow water while slowly walking towards the main water source, thus stepping steadily over the benthonic mat over which multiple fish were swimmin

A theropod trackway providing evidence of a pathological foot from the exceptional locality of Las Hoyas (upper Barremian, Serranía de Cuenca, Spain)

We describe a trackway (LH-Mg-10-16) occurring in laminated carbonated limestones of the Las Hoyas locality, Serranía de Cuenca, Spain. It is unmistakably a large theropod dinosaur trackway encompassing two unusual aspects, namely, wide-steps, and a set of equally deformed left footprints (with a dislocated digit). The layer also preserves other vertebrate trails (fish Undichna) and different impressions in the sediment. To address these complex settings, we devised a multidisciplinary approach, including the ichnological and taphonomical descriptions, characterisation of the rock lithofacies using thin-sections, 3D structured-light digitalisation with a high precision of 200-400 μm, and a geometric morphometric comparison with a large sample of bipedal dinosaur trackways. Sedimentary analyses showed that the trackway was produced in a humid, benthonic microbial mat, the consistency and plasticity of which enabled the preservation of the details of the movement of the animal. The results of the geometric analysis indicate that the 'wide-steps' of the trackway is not unusual compared to other trackways, providing evidence that it was made by a single individual with an estimated hip height approximately 2 m. Analogous pathologies in extant archosaurs that yield the combination of wide steps and deformed digits in the same trackway were considered. All results mutually support the hypothesis that a large theropod dinosaur, with a pathological foot, generated the trackway as it crossed an area of shallow water while slowly walking towards the main water source, thus stepping steadily over the benthonic mat over which multiple fish were swimming

Table S1 from Open data and digital morphology

Summary of main online repositories for 3D digital morphological data