Archaeopteryx lithographica, specimen displayed at the Museum für Naturkunde in Berlin. (From Wikimedia Commons)
Birds originated from a theropod lineage more than 150 million years ago. Their evolutionary history is one of the most enduring and fascinating debates in paleontology. In recent years, several discovered fossils of theropods and early birds have filled the morphological, functional, and temporal gaps along the line to modern birds. The discovered fossils demonstrate that distinctive bird characteristics such as feathers, flight, endothermic physiology, unique strategies for reproduction and growth, and a novel pulmonary system have a sequential and stepwise transformational pattern, with many arising early in dinosaur evolution, like the unusually crouched hindlimb for bipedal locomotion,the furcula and the “semilunate” carpal that appeared early in the theropod lineage (Allen et al., 2013; Xu et al., 2014). Also, the discovery of Mahakala – a basal dromaeosaurid dinosaur named for one of the eight protector deities in Tibetan Buddhism – suggests that extreme miniaturization and laterally movable arms necessary for flapping flight are ancestral for paravian theropods. In contrast, a number of basal birds resemble theropods in many features.
Sciurumimus (A); the basal coelurosaur Sinosauropteryx (B) with filamentous feathers; the deinonychosaurs Anchiornis (C) and Microraptor (D). Adapted from Xu et al., 2014.
Anatomical features like aspects of egg shape, ornamentation, microstructure, and porosity of living birds trace their origin to the maniraptoran theropods, such as oviraptorosaurs and troodontids. In addition, some preserving brooding postures, are known for four oviraptorosaurs, two troodontids, a dromaeosaur, and one basal bird providing clear evidence for parental care of eggs.
In birds, particularly their forebrains, are expanded relative to body size. The volumetric expansion of the avian endocranium began relatively early in theropod evolution. Archaeopteryx lithographica is volumetrically intermediate between those of more basal theropods and crown birds (Balanoff et al., 2013). The digital brain cast of Archaeopteryx also present an indentation that could be from the wulst, a neurological structure present in living birds used in information processing and motor control with two primary inputs: somatosensory and visual. Birds also exhibit the most advanced vertebrate visual system, with a highly developed ability to distinguish colors over a wide range of wavelengths.
Reconstruction of pulmonary components [cervical air-sac system (green), lung (orange), and abdominal air-sac system (blue)] in the theropod Majungatholus (From Xu et al., 2014)
Feathers were once considered to be unique avialan structures. The megalosaurus Sciurumimus
, the compsognathus Sinosauropteryx,
and a few other dinosaurs, document the appearance of primitive feathers. More recent studies indicated that non avian dinosaurs, as part of Archosauria, possessed the entirety of the known non keratin protein-coding toolkit for making feathers (Lowe et al., 2015)
The evolution of flight involved a series of adaptive changes at the morphological and molecular levels,like the fusion and elimination of some bones and the pneumatization of the remaining ones. The extensive skeletal pneumaticity in theropods such as Majungasaurus demonstrates that a complex air-sac system and birdlike respiration evolved in birds’ theropod ancestors. The increased metabolism associated with homeothermy and powered flight requires an efficient gas exchange process during pulmonary ventilation. Moreover, recent anatomical and physiological studies show that alligators, and monitor lizards exhibit respiratory systems and unidirectional breathing akin to those of birds, which indicate that unidirectional breathing is a primitive characteristic of archosaurs or an even more inclusive group with the complex air-sac system evolving later within Archosauria.
The earliest diversification of extant birds (Neornithes) occurred during the Cretaceous period and after the mass extinction event at the Cretaceous-Paleogene (K-Pg) boundary, the Neoaves, the most diverse avian clade, suffered a rapid global expansion and radiation. Today, with more than 10500 living species, birds are the most species-rich class of tetrapod vertebrates.
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