On the morphological description of tracheal and esophageal displacement and its phylogenetic distribution in avialae

Abstract

This research examines the evolution and phylogenetic distribution of a peculiar and often overlooked character seen in birds, herein called tracheal and esophageal displacement. Tracheal and esophageal displacement refers to an asymmetrically situated trachea and/or esophagus along the length of the neck. This contrasts with what would be perceived as the "normal" (midsagittal) placement of these organs, wherein the two organs are situated along the ventral midline of the neck with no deviation. A total of forty-two bird species were examined (thirty-six of which came from dissections whereas six came from comments from previous literature or personal observations), as well as turtles, lizards, crocodylians, and mammals. This study found that essentially all birds have a laterally displaced trachea and/or esophagus. Lizards and mammals were seen to have normal, midsagittally placed tracheae and esophagi. Crocodylians were interesting in that alligators were defined by a normally situated trachea and esophagus whereas some crocodiles were characterized by displacement. In birds, the displacement may occur gradually along the neck, or it may happen immediately upon exiting the oropharynx. Displacement of these organs in birds is the result of a heavily modified neck wherein muscles that restrict mobility of the trachea and esophagus in lizards, alligators, and mammals (e.g., m. episternocleidomastoideus, m. omohyoideus, and m. sternohyoideus) no longer substantially restrict positions of the trachea and esophagus in birds. Rather, these muscles are modified in ways which may assist with making tracheal movements. The implications of this study may provide interesting insights for future comparisons in extinct taxa.