Interplay of size and shape in miniaturized land snails
ABSTRACTThe distribution of the mean greater shell dimensions of 2446 genera of stylommatophoran land snails consists of two groups peaking at 3 mm and 15.3 mm. The 3-mm group includes the miniature snails whose adult shell dimensions range from almost 0.8 mm to less than 1.5 mm. Relative surface area, shell thickness and egg size are discussed as potential factors that may limit the minimum shell dimensions. To obtain uniform distributions, the shell shape is expressed as the expanse angle, defined as the apex angle of a right triangle the height and the base of which are the shell height and diameter, respectively. In terms of expanse angles there are three shell shapes: tall (< 40°), flat (> 50°) and roughly equiaxial (40°-50°). The variation of shell shape with size was analyzed within the morphospace they form. At shell dimensions above 10 mm the lowest expanse angle is about 10°. Below 10 mm, the lower limit of the expanse angle increases as shells get smaller. As a result, no miniature species has a tall shell. It is proposed that two evolutionary functional constraints render small and narrow shells of miniaturized snails nonadaptive. These are the requirements to reduce surface areas to decrease water loss and to have enough volume and a wide enough body whorl to accommodate at least one egg.