Climate warming is discussed as a factor that can favour the success of invasive species. In the present study, we analysed potential fitness gains of moderate warming (3 1C above field temperature) on the invasive clam Corbicula fluminea during summer and winter. The experiments were conducted under seminatural conditions in a bypasssystem of a large river (Rhine, Germany). We showed that warming in late summer results in a significant decrease in the clams' growth rates (body mass and shell length increase) and an increase in mortality rate. The addition of planktonic food dampens the negative effect of warming on the growth rates. This suggests that the reason for the negative growth effect of temperature increase in late summer is a negative energetic balance caused by an enhanced metabolic rate at limited food levels. Warming during early summer revealed contrasting effects with respect of body mass (no warming effect) and shell length (increased shell growth with warming). This differential control of both parameters further enhances the loss of the relative (size-specific) body mass with warming. In contrast, warming in winter had a consistently positive effect on the clams' growth rate as demonstrated in two independent experiments. Furthermore, the reproduction success (as measured by the average number of larvae per clam) during the main breeding period (April) was strongly enhanced by experimental warming during winter, i.e. by eight times during the relatively cold winter 2005/2006 and by 2.6 times during the relatively warm winter 2007/2008. This strong, positive effect of moderate winter warming on the clams' fitness is probably one reason for the recent invasion success of C. fluminea in the northern hemisphere. However, warm summer events might counteract the positive winter warming effect, which could balance out the fitness gains.

The filter-feeding clam Corbicula fluminea has widely spread from its Asiatic origin into freshwater habitats on several continents, where it often has a considerable impact on ecosystem processes. The present study was stimulated by the observation that C. fluminea can experience mass mortality during warm summers, even when temperatures are still far below the lethal level. We hypothesized that starvation due to low food quantities during summer is a main factor in this context. In order to test nutritional conditions in an environment where summer mortality occurred, the clams' body mass was tracked in river bypass systems installed at the Lower River Rhine (Germany and The Netherlands). Two food levels were adjusted in the bypass channels: one corresponding to the original chlorophyll level in the river (ambient food) and one with a chlorophyll a level reduced by about 50% (low food). The clams kept at the ambient food level increased their shell length during summer, although growth rates decreased at low food levels in the River Rhine in late summer. In contrast to shell length, body mass decrease in late summer cumulated in 94% reduction from August until October. This trend was enhanced by the experimental food reduction, i.e., clams kept in the low food level treatments weighed 60% less than the clams in the ambient food treatment at the end of summer. However, mortality was low in both treatments. The data demonstrate a high plasticity in the body mass of C. fluminea. The corresponding possibility to respire body mass can be seen as one strategy to survive longer starvation periods.

1. Both the clearance rates (CR) and abundances of the freshwater sponge Ephydatia muelleri and the bryozoans Plumatella emarginata and Fredericella sultana were investigated from autumn to spring under different temperature regimes. The experiments were performed in bypass channels of the River Rhine (Cologne, Germany) in which temperature could be manipulated. 2. The impact of temperature increase on CRs depends upon the grazer: E. muelleri showed a clear increase in CRs with increasing temperature whereas P. emarginata was not significantly affected by experimental warming. 3. Distinct differences in food preference were found for the sponge (which is an efficient grazer of bacteria and small algae) and for the bryozoan P. emarginata (which feeds primarily on large algae, and with no significant grazing on bacteria). 4. In contrast to their temperature-related patterns in CR, respiration of both P. emarginata and E. muelleri increased with temperature between 19 and 32°C, suggesting that the risk of experiencing energy deficiency at high temperatures due to increased metabolic rates is particularly high for the bryozoan. 5. A temperature elevation of 3°C above the natural Rhine temperature resulted in a delay in the disappearance of active tissue and formation of resting stages for E. muelleri in autumn. This delay ranged from 8 (beginning of gemmulation) to 22 days (termination of gemmulation). In contrast, there was no distinct effect of warming on the disappearance of active zooids of the two bryozoan species in autumn. However, warming can positively affect the maintenance of active zooids during winter in F. sultana. In spring, the appearance of active zooids of P. emarginata was clearly stimulated by temperature elevations, whereas the hatching of both F. sultana and E. muelleri was hardly affected by warming. 6. The study demonstrated different patterns in the thermal ecology of the two freshwater bryozoans and the sponge in comparison to other filter feeders, particularly mussels. Such patterns need to be considered when predicting the impact of temperature on pelagic-benthic coupling in aquatic habitats.