Systematics and geographical distribution of Galba species, a group of cryptic and worldwide freshwater snails

Abstract

Cryptic species can present a significant challenge to the application of systematic and biogeographic principles, especially if they are invasive or transmit parasites or pathogens. Detecting cryptic species requires a pluralistic approach in which molecular markers facilitate the detection of coherent taxonomic units that can then be analyzed using various traits (e.g., internal morphology) and crosses. In asexual or self-fertilizing species, the latter criteria are of limited use. We studied a group of cryptic freshwater snails (genus Galba) from the family Lymnaeidae that have invaded almost all continents, reproducing mainly by self-fertilization and transmitting liver flukes to humans and livestock. We aim to clarify the systematics, distribution and phylogenetic relationships of these species with an integrative approach that includes morphology (shell and reproductive anatomy), molecular markers, wide-scale sampling across America, and data retrieved from GenBank (to include Old World samples). Our phylogenetic analysis suggests that the genus Galba originated ca. 22 Myr ago and today comprises six clusters of species. Four of these clusters (G. truncatula, G. cubensis/viator, G. humilis and G. schirazensis) are morphologically cryptic and constitute species or species complexes with wide variation in their genetic diversity, geographic distribution and invasiveness. The other two clusters constitute a single species or a species complex (Galba cousini/meridensis) that demonstrate more geographically restricted distributions and exhibit an alternative morphology more phylogenetically derived than the cryptic one. Further genetic studies are required to clarify the status of both G. cousini/meridensis and G. cubensis/viator. We emphasize that no Galba species should be identified without molecular markers and that additional sampling is required, especially in North America, Eurasia and Africa to clarify remaining questions in systematics and biogeography. We also discuss several hypotheses that can explain crypsis in Galba, such as convergence and morphological stasis, and hypothesize a role for stabilizing selection in amphibious (rather than strictly freshwater) habitats.