Dikerogammarus villosus, a Ponto-Caspian species, is known to be a particularly successful invader, and is currently the prevailing invasive gammarid in Southern German large waters. Field observations suggest that D. villosus has replaced the native G. pulex and the invasive D. haemobaphes, also a Ponto-Caspian species, in some reaches of the German Danube. Dikerogammarus villosus is also believed to be the reason why Echinogammarus berilloni, a Mediterranean species, and Dikerogammarus bispinosus, a further Ponto-Caspian gammaridean species, could not build significant populations in Southern German rivers. Because intraguild predation (IGP) is regarded as a major force leading to species displacements, we hypothesized that superiority in predation by D. villosus is responsible for the disappearance or limited occurrence of several native and invasive gammarideans in many reaches of Southern German waters. To test this hypothesis, we conducted a series of laboratory IGP experiments with 1:1 combinations in which D. villosus was allowed to prey on other gammarids and vice versa. We also studied the extent of cannibalism within species. Dikerogammarus villosus was a stronger predator than G. pulex and E. berilloni. However, D. villosus was not stronger in predation than its relatives D. haemobaphes and D. bispinosus, although results with the combinations of D. villosus and D. bispinosus were less conclusive, especially in the female-female combination. Cannibalism rate was high in D. haemobaphes compared with other species. We conclude that superiority in IGP experiments could explain the disappearance of G. pulex and the missing or limited occurrence of E. berilloni since the arrival of D. villosus in Southern German rivers. However, the results of IGP experiments do not explain why D. haemobaphes was displaced by D. villosus in many places and why D. bispinosus could not build significant populations in Southern German waters. Possibly specific combinations between mutual predation and cannibalism or factors other than predation could have affected displacement and occurrence patterns of Dikerogammarus species in Southern German waters.
Substrate choice, swimming activity and risk to predation by burbot (Lota lota) of the well established Gammarus roeselii and the invader Dikerogammarus villosus were studied in mixed and single-species aquarium experiments. We used stones, gravel and aquatic weeds (Elodea, Chara) as substrates. We hypothesized that both species have different substrate preferences and that substrate affects the predation risk. We also assumed that presence of D. villosus influences substrate preference and predation risk of G. roeselii since the invader is known to affect the behavior of other gammarids. Adults of D. villosus in single species experiments and juveniles in mixed and single species experiments were evenly distributed over the different substrates but adults in mixed species experiments were more likely to prefer stone substrate. In contrast, adults and juveniles of G. roeselii clearly preferred aquatic weeds independent of the presence/absence of the invader. Both species preferred substrates with fissured surface over substrates with smooth surface. Gammarus roeselii was observed swimming more often than D. villosus in the open water but its swimming activity was lower when its preferred substrate was present compared with its swimming activity if non-preferred substrates were present. Predation rate of burbot on D. villosus was comparatively low and independent of the substrate. Burbot consumed many more G. roeselii than D. villosus, both in mixed and single species experiments. But when the preferred substrate of G. roeselii (weeds) was used in the experiments, predation rate of burbot on G. roeselii was somewhat lower than that when non-preferred substrates were present. The results of the experiments support our hypothesis that the gammarids studied here have different substrate preferences and that presence of the preferred substrate can affect predation risk. However, there is no evidence that presence of D. villosus affected substrate choice or predation risk in G. roeselii. We consider that differences in use of spatial niches permit co-existence of G. roeselii and D. villosus in the wild when substrates are diverse. The fact that G. roeselii than D. villosus is more often observed swimming in the open water may explain its higher risk of being captured by fish.
The gammarid composition at 25 sites in the rivers Danube, Main and the southern reaches of the Rhine were studied during the years 2002-2004. Dikerogammarus villosus and Echinogammarus ischnus were the most frequent species prevailing at 17 sites. Sympatric occurrence of D. villosus and E. ischnus was observed at 12 sites. Dikerogammarus haemobaphes was recorded at 7 sites; this species prevailed in the Danube, west of the Weltenburger Enge and in the Isar mouth where it co-existed with native species (Gammarus pulex and/or G. roeseli) at 6 sites. Dikerogammarus bispinosus and E. berilloni were found at only 1 site, where they co-existed with D. villosus and E. ischnus, and with D. villosus and native species, respectively. Investigation of reproductive characteristics at 3 sites showed that females of D. villosus and D. haemobaphes produced the biggest clutches with more than 100 eggs. Females of E. ischnus produced much smaller clutches (10-35 eggs on an average), but very big eggs. Clutch sizes and egg volumes of D. bispinosus and E. berilloni resembled those of native species. Our results suggest that the most successful invaders (D. villosus, D. haemobaphes and E. ischnus) display reproductive traits that facilitate their success. Both Dikerogammarus sp. allocate energy into production of many but small eggs, thus maximizing offspring number, while E. ischnus allocates its energy into production of fewer but large eggs which could be beneficial at sites where food is scarce.
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