The proboscis of tapirs (Mammalia: Perissodactyla): a case study in novel narial anatomy

The trunk-like proboscis of tapirs provides a prime case study in the evolution of anatomical novelty. Morphological study of this unique structure was undertaken employing several specimens and a combination of analytical techniques: gross anatomical dissection, radiographic imaging and histological sectioning. Evolution of the proboscis of tapirs entailed wholesale transformation of the narial apparatus and facial architecture relative to perissodactyl outgroups. This transformation involved retraction and reduction of the bony and cartilaginous facial skeleton, such that several structures present in outgroups are completely absent in tapirs, including cartilages surrounding the nasal vestibule (e.g. alar and medial accessory cartilages, rostral portion of the nasal septum) and associated musculature (dilatator naris apicalis, lateralis nasi pars ventralis). At the same time, soft tissues surrounding the upper lip and nose became elaborated to form a mobile, ̄eshy proboscis. Several key facial muscles (e.g. levator labii superioris, levator nasolabialis, caninus, lateralis nasi) have been co-opted to function in movement of the proboscis. The nasal vestibule is expanded and occupies approximately 75% of the nasal cavity. Vestibular expansion has compressed and simpli®ed caudal components of the nasal cavity (e.g. reduction of dorsal and middle nasal conchae, loss of plica recta and plica basalis). The airway has become dorsally arched causing the ventral conchal complex to become inclined relative to the long axis of the skull. A few anatomical enigmas remain, such as the complicated maxilloturbinate that rostrally contacts the nasal septum and vomeronasal organ. Similarly, the meatal diverticulum, despite being both ancient and anatomically complex, has no obvious functional signi®cance; it is clear that it is not homologous to the nasal diverticulum of horses and other equids. The reduction of the osseocartilaginous portion of the proboscis, coupled with expansion of the muscular and connective tissue components, has resulted in an organ that is best interpreted as a muscular hydrostat.