The evolution of chemoreception in squamate reptiles: a phylogenetic approach

Recent advances in the field of squamate reptile chemoreception have been paralleled by the growth and preeminence of cladistics in the field of systematics, but for the most part, workers in the former have failed to incorporate the conceptual and informational advances of the latter. In this paper, I attempt a preliminary rapprochement by combining the methods of phylogenetic systematics and current hypotheses of squamate relationships with an overview of squamate chemosensory biology. This purely phylogenetic approach leads to a number of falsifiable generalizations about the evolution of chemoreception in squamates: 1) Evolution of this system is conservative rather than plastic, reflecting to a large extent suprafamilial attributes rather than adaptation to local conditions; 2) Anguimorphs are highly chemosensory and teiids show convergence with this group; 3) Tongue-flicking, a bifurcated tongue tip, a vomeronasal (VNO) mushroom body, and a complete circular muscle system in the tongue are a correlated character complex associated with the attainment, in squamates, of a direct VNO-oral connection and the loss of a VNO-nasal connection; 4) There is little support for a visual-chemosensory dichotomy within Squamata; 5) Gekkotans are allied with Autarchoglossa, both phylogenetically and in terms of chemosensory biology; 6) Iguania are highly variable in chemosensory development; iguanids represent the primitive iguanian condition, while agamids and chamaeleonids have secondarily reduced or lost their chemosensory abilities; 7) Apparent contradictions in chemosensory behavior among iguanids probably represent intrafamilial divergence; 8) Ecological correlates within Iguanidae and other taxa might be spurious, resulting from historical factors unrelated to the adaptations in question; 9) The mechanical demands of lingual food prehension have constrained chemosensory evolution in Iguania; chemosensory evolution within Scleroglossa was permitted by the liberation of the tongue from this ancestral role.