Environment, intraspecific lineages and geographic range jointly shape the high morphological variability of Dikerogammarus villosus (Sowinsky, 1894) (Crustacea, Amphipoda): a successful aquatic invader across Europe

Phenotypic variability is a key factor promoting the establishment and spread of invasive populations in new environments. The Ponto-Caspian region contains a diverse endemic fauna known for its exceptional environmental plasticity, with many species invading European waters. However, the extent to which the environment shapes the phenotypic variability of these successful invaders remains poorly understood. We test to what extent the environment, intraspecific lineage affinity and geographic range interact and shape the variability of ecologically relevant functional morphological traits of the amphipod, Dikerogammarus villosus. Our results show the highest differentiation among environments, with an enhancement of predation-related traits in brackish waters relative to freshwaters. Differentiation among lineages and ranges (native/invaded) was smaller, occurring in traits related to locomotion and food processing. Although we uncovered an overall increase in variability outside the native range, the dynamics of morphological change were lineage-specific: the Western Lineage (invading via the River Danube) underwent a shift towards increased appendage length, while the Eastern Lineage (invading via the River Dnieper) underwent a significant overall morphospace expansion. We conclude that D. villosus exhibits a remarkable morphological variability across Europe that is influenced by the interplay between the environment as well as its evolutionary and invasion history.