Integration of pecking, filter feeding and drinking mechanisms in waterfowl

Abstract

This paper is one of several contributions in a series, illustrating the application of a specific deductive methodology to explain diversity of form. The methodology facilitates the explanation of feeding morphologies in various ducks as a transformation of the mallard's feeding design maximized for specific proportions of performance that are contributed by pecking and filter feeding mechanisms.

The earlier described anatomy and formal analyses of the three mechanisms in the mallard served as the initial conditions used in simulation models. Four elements of the feeding system were chosen that play a major role in all three mechanisms. For each element, the main parameter was selected: storage capacity of the rostral mouth cavity, transport capacity of the rostral mouth tube, storage capacity of the caudal mouth cavity and transport capacity of the caudal mouth tube. The boundary conditions for the simulation were determined from internal organismic constraints. The total food uptake of the mallard was regarded as the function to be maximized. This ‘object’ function is the summation of the food uptake by one second of pecking and one second of filter feeding. The drinking mechanism was shown not to interfere, since that mechanism operates sufficiently whenever the pumping mechanism works properly.

The ‘object’ function, made up by the pecking and filter feeding performances was graphed. From these graphs a morphospace was developed: the region within which modifications of the mouth design are feasible. This procedure allowed examination of the general hypothesis that different modifications of one design for a complex multi-role system are explainable from differences in proportions of the functional performance contributed by each of the roles.

Two predictions were evaluated more specifically: 1) If filter feeding performance must increase for a specific change in total food uptake, the volume of the rostral mouth cavity must increase; this requires widening and lengthening of the rostral maxillar portion and also a phase shift in jaw and lingual motion patterns, increasing the stroke volume. 2) If pecking performance must increase, the transport capacity of the rostral mouth tube must increase; this requires shortening of the maxillar mid portion. These two predictions regarding change in mouth morphology were borne out by shovelers and tufted ducks, respectively.

The deductive analysis of the feeding system's morphospace allowed the explanation of the wide diversity of forms in the duck mouth designs as emerging from the great radiation capacity of the ancestral duck's mouth design: it is a versatile, potentially multi-role system in which pecking, drinking and filter feeding mechanisms are mutually very tolerant and, by epiphenomena, includes the elements of a grazing mechanism. In addition, the deduced morphospace was shown to explain phenotypic plasticity, since it explains the occurrence of two morphotypes that develop due to habitat discrimination: one type that has been forced to filter feed, and one type that feeds by pecking.