Winning the biodiversity arms race among freshwater gastropods: competition and coexistence through shell variability and predator avoidance

Explanations for the coexistence of many closely related species in inland waters continue to be generated more than 50 years after Hutchinson's question: why are there so many kinds of animals? This review focuses on the hypothesis that high species diversity of freshwater gastropods results, in part, from predators maintaining biodiversity across a range of deep- and shallow-water habitats. Invertebrate predators, such as aquatic insects, and leeches consume soft tissue of pulmonate snails by penetrating shells of various shapes and sizes. Crayfish and large prawns chip around the shell aperture to enter thick shells and crush small shells with their mandibles. Crabs use their strong chelae to crush thin and thick shells. Fishes with pharyngeal teeth are major shell-breaking predators that combine with other vertebrate predators such as turtles and wading birds to increase the diversity of gastropod communities by regulating the abundance of dominant species. Although the generalized diets of most freshwater predators preclude tight co-evolutionary patterns of responses, there are combinations of predators that modify gastropod behavior and shell morphology in aquatic assemblages of different ages and depths. This combination of invertebrate and vertebrate predatory impacts led to competitive advantages among individual gastropods with different adaptations: (1) less vulnerable shell morphologies and sizes; (2) predator-avoidance behaviors; or (3) rapid and widespread dispersal with variable life histories. Some individuals develop thicker and/or narrow-opening shells or shells with spines and ridges. Other thin-shelled species crawl out of the water or burrow to lower their risk to shell-breaking or shell-entering predators. Some alter their age at first reproduction and grow rapidly into a size refuge. Fluctuations in water levels and introductions of non-native species can change competitive dominance relationships among gastropods and result in major losses of native species. Many different gastropod predators control species that are human disease vectors. Most snails and their predators provide other ecosystem services such as nutrient cycling and transfer of energy to higher trophic levels. Their persistence and diversity of native species require adaptive management and coordinated study.