The most complete known North American enantiornithine was collected in 1992 but never formally described. The so-called “Kaiparowits avisaurid” remains one of the most exceptional Late Cretaceous enantiornithine fossils. We recognize this specimen as a new taxon, Mirarce eatoni (gen. et sp. nov.), and provide a complete anatomical description. We maintain that the specimen is referable to the Avisauridae, a clade previously only known in North America from isolated tarsometatarsi. Information from this specimen helps to clarify evolutionary trends within the Enantiornithes. Its large body size supports previously observed trends toward larger body mass in the Late Cretaceous. However, trends toward increased fusion of compound elements across the clade as a whole are weak compared to the Ornithuromorpha. The new specimen reveals for the first time the presence of remige papillae in the enantiornithines, indicating this feature was evolved in parallel to dromaeosaurids and derived ornithuromorphs. Although morphology of the pygostyle and (to a lesser degree) the coracoid and manus appear to remain fairly static during the 65 million years plus of enantiornithine evolution, by the end of the Mesozoic at least some enantiornithine birds had evolved several features convergent with the Neornithes including a deeply keeled sternum, a narrow furcula with a short hypocleidium, and ulnar quill knobs—all features that indicate refinement of the flight apparatus and increased aerial abilities. We conduct the first cladistic analysis to include all purported avisuarid enantiornithines, recovering an Avisauridae consisting of a dichotomy between North and South American taxa. Based on morphological observations and supported by cladistic analysis, we demonstrate Avisaurus to be paraphyletic and erect a new genus for “A. gloriae,” Gettyia gen. nov.
In recent years, the Lower Cretaceous (Aptian) Xiagou Formation has yielded approximately 100 avian partial skeletons, many with soft-tissue traces, from sites in the Changma Basin of Gansu Province, north-western China. The most abundant taxon amongst these is the ornithuromorph Gansus yumenensis, but enantiornithines have also been identified in the sample. Here we describe two incomplete, semi-articulated appendicular skeletons, the first consisting of a partial left pelvic girdle and complete pelvic limb, and the second comprised of a nearly complete right pelvic limb. Both specimens bear characteristics diagnostic of Enantiornithes, and are referred to a new taxon, Qiliania graffini gen. et sp. nov. The exceptional, three-dimensional preservation of these specimens (compared to the crushed, nearly two-dimensional condition of most other Early Cretaceous avian fossils) reveals new information regarding enantiornithine anatomy, evolution, and diversity.
Finely−bedded lacustrine deposits of the Aptian (Lower Cretaceous) Xiagou Formation exposed in the
Most crown-birds experience rapid growth, reaching adult size within a year. Rapid growth strategies evolved within Aves multiple times during the Cretaceous, documented in the Confuciusornithiformes and the Ornithuromorpha. In contrast, osteohistological data suggest the Enantiornithes, the dominant clade of Cretaceous terrestrial birds, were characterized by much slower growth rates that were sustained longer into adulthood. Here we provide evidence for a unique growth strategy involving relatively rapid growth in the Late Cretaceous avisaurid enantiornithine, Mirarce eatoni. Multiple appendicular skeletal elements were sectioned for osteohistological analysis. These show remarkable intraskeletal variation, and high levels of variation even between individual sections. The radius is composed of parallel-fibered bone, similar to histological descriptions in other enantiornithines. Other elements, in contrast, differ markedly from other members of the clade. The humerus is composed of parallel-fibered bone with a middle layer of incipient fibrolamellar bone and several growth lines in the outer circumferential layer and near the endosteal border. The endosteal and periosteal layers of slow-growing bone indicate cyclical variation in growth rates. The femur shows regions of coarse compact cancellous bone and parallel-fibered bone with numerous secondary osteons, and only a single growth line. The tarsometatarsus is predominantly fibrolamellar in texture, with several asymmetrical growth lines located throughout the cortex; this element exhibits strong cortical drift. Growth lines in both the endosteal and periosteal portions of the cortex indicate that, like the humerus, growth rates of this bone varied cyclically. The two phalanges studied here are composed of parallel-fibered bone with extensive evidence of and remodeling over possible regions of coarse compact cancellous bone. Although Mirarce is one of the largest known enantiornithines, slow and protracted growth documented in similarly-sized taxa suggests this bone texture is not merely a size-related scaling effect. These findings indicate that by the Late Cretaceous, some enantiornithines had evolved absolutely higher growth rates and more complex life history strategies, in which growth rates varied across the skeleton. Furthermore, a variety of strategies were employed to achieve adult size and morphology, including cycles of slow and fast growth, asymmetrical growth within a single element, and extensive remodeling.
The relative length proportions of the three bony elements of the pelvic (femur, tibiotarsus and tarsometatarsus) and pectoral (humerus, ulna and manus) limbs of the early Cretaceous bird Gansus yumenensis, a well-represented basal ornithuromorph from China, are investigated and compared to those of extant taxa. Ternary plots show that the pectoral limb length proportions of Gansus are most similar to Apodiformes (swifts and hummingbirds), which plot away from all other extant birds. In contrast, the pelvic limb length proportions of Gansus fall within the extant bird cluster and show similarities with the neornithine families Podicipedidae (grebes), Diomedeidae (albatross) and Phalacrocoracidae (cormorants). Although it does have some of the pelvic limb features of grebes and cormorants, the femur of Gansus is more gracile and is thus more consistent with an albatross-like shallow-diving mode of life than a strong foot-propelled diving movement pattern. The position of Gansus in pectoral limb ternary morphospace is largely due to its elongated manus. In contrast to apodiformes, where the humerus and ulna are short and robust, an adaptation, which provides a stiff wing for their demanding fast agile and hovering flight (respectively), the wing-bones of Gansus are slender, indicating a less vigorous flapping flight style. The suite of characters exhibited by Gansus mean it is difficult to completely interpret its likely ecology. Nevertheless, our analyses suggest that it is probable that this bird was both volant and capable of diving to some degree using either foot-propelled or, perhaps, both its wings and its feet for underwater locomotion.
Bone histology of crown-group birds is a research topic of great interest, permitting insight into the evolution of remarkably high growth rates in this clade and variation across the altricial-precocial spectrum. In this study, we describe microanatomical characteristics of the humerus and femur in partial growth series from 14 crown group birds representing ten major clades (Struthioniformes, Galliformes, Apodiformes, Columbiformes, Charadriiformes, Accipitriformes, Strigiformes, Psittaciformes, Falconiformes, and Passeriformes). Our goals were to: (1) describe the microanatomy of each individual; (2) make inter-and intra-taxonomic comparisons; (3) assess patterns that correspond with developmental mode; and (4) to further parse out phylogenetic, developmental, and functional constraints on avian osteological development. Across taxa, the femoral and humeral tissue of neonates can be broadly characterized as highly-vascularized, disorganized woven bone with great variation in cortical thickness (inter-and intrataxonomically, within an individual specimen, and within a single section). The tissue of precocial chicks is relatively more mature at hatching than in altricial, but other categories along the developmental spectrum were less easy to distinguish, thus we were unable to identify a definitive histological proxy for developmental mode. We did not find evidence to support hypotheses that precocial chicks exclusively have thicker cortices and more mature bone in the femur than the humerus at time of hatching; instead, this is a characteristic of nearly all taxa (regardless of developmental mode), suggesting deep evolutionary origins and the effects of developmental channeling. Bone tissue in adults exhibited unexpected variation, corresponding to differences in body size. Large-bodied birds have cortices of fibrolamellar bone, but organization of tissue increases and vascularity decreases with diminishing body size. The outer circumferential layer (OCL) also appears at earlier growth stages in small-bodied taxa. Thus, while the OCL is indicative of a cessation of appositional growth it is not always indicative of cortical maturity (that is, maximum organization of bony tissue for a given taxon). Small size is achieved by truncating the period of fast growth; manipulation of the timing of offset of bone growth is therefore an important factor in changing growth trajectories to alter adult body size.
A partial skeleton of a sauropod dinosaur, Museum of Western Colorado 8028, was recovered from the Upper Jurassic Morrison Formation near Snowmass, Colorado, USA, and in 2014 it was referred to Haplocanthosaurus sp. The material includes four proximal caudal vertebrae, which are unique in having strongly amphicoelous vertebral bodies, in which the centrum is reduced to a thin vertical plate of bone between the concave articular surfaces, and neural canals that are laterally and ventrally expanded and strongly sloped relative to the centrum. To reconstruct the soft tissues that left these traces on the skeleton, we rebuilt these vertebrae using both physical and digital techniques. We CT scanned the specimens, generated digital models, 3D-printed the vertebrae, physically sculpted missing material onto the printed models, optically scanned the sculpted vertebrae to create second-generation digital models, and finally, retro-deformed and articulated those digital versions of the vertebrae. The spaces between the deeply amphicoelous caudal centra were likely filled by large, ellipsoidal intervertebral discs, as in the amphicoelous vertebrae of Sphenodon and gekkotan lizards. The expanded neural canals remain enigmatic. In ostriches, the lumbosacral spinal cord is expanded laterally and ventrally in each vertebra, lending the spinal cord a shape that roughly resembles beads on a string. Similar spinal cord morphology might explain the expanded neural canals in the Snowmass Haplocanthosaurus, but it is not clear why a relatively small-bodied, small-tailed sauropod would need such a spinal expansion, given that similar expansions have not been reported in larger-bodied and larger-tailed taxa.
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