Conroy is correct to note that our experimental study did not take into account how hydrodynamic coupling affected the ability of zebra mussels to graze algae. Unfortunately, the size of our mesocosms (18 L) did not allow for, nor were they designed to provide, a thorough quantification of benthic–pelagic coupling. Therefore, we agree with Conroy that the magnitude of zebra mussel effects on phytoplankton within individual systems will depend on site-specific factors such as turbulent mixing, mussel filtering efficiency, phytoplankton reproduction and sinking rates, and nutrient remineralization. As such, we made no attempts to directly quantify zebra mussel filtration or grazing rates in our study reservoir. Instead, we presented our research as “a preliminary study” and suggest that additional research is needed to address among other things “ . . . how zebra mussels affect algal responses to nutrient enrichment in experiments that better mimic natural conditions (e.g. large in situ reservoir enclosures).” With that said, however, we do not agree with Conroy when he states that our proposal is inconsistent with published examples of post-invasion increases in phytoplankton, benthic algae and cyanobacterial toxins. For example, he provides two references (Raikow et al., 2004; Conroy et al., 2005) to show that phytoplankton have increased following invasion. However, as cited in our original manuscript, there are also numerous examples of phytoplankton decreasing following invasion; even Raikow et al. (Raikow et al., 2004) reported that lakes with zebra mussels had lower water column concentrations of total phytoplankton biomass than lakes without zebra mussels. Furthermore, zebra mussels can “decouple” relationships between total phosphorus (TP) and chlorophyll a so that observed yields of algal biomass are less than predicted from TP concentrations (see Nichols et al., 2001 and other references cited in Dzialowski and Jessie, 2009). It is also important to note that we address how zebra mussels may impact both benthic algae (e.g. they may increase) and cyanobacteria (e.g. the effects may be nutrient dependent) in eutrophic reservoirs in light of recently published research [see Dzialowski and Jessie, 2009 for more details]. Combined, we believe that these studies are consistent with our predictions that “ . . . invaded reservoirs may exhibit lower yields of algal biomass than those predicted from TP concentrations . . . ” and that “ . . . the severity of algal bloom events may be lower in invaded reservoirs if zebra mussels are able to directly negate or mask the increasing effects of external nutrient pulses on algal biomass as we have shown here.” We would also like to respond to Conroy’s assertion that the effects of zebra mussels on phytoplankton are most apparent in the 1 m boundary layer at the bottom of lakes. While this appears to be true for the western basin of Lake Erie, it is also important to note that zebra mussels have reduced phytoplankton biomass in the epilimnion of stratified lakes and reservoirs as well (Yu and Culver, 2000; Raikow et al., 2004; Knoll et al., 2008). Therefore, it seems inappropriate to suggest that the effects of zebra mussels on phytoplankton will be restricted mainly to the benthos. As such, additional research is needed to determine how hydrodynamic coupling affects the ability of zebra mussels to impact systems other than the Great Lakes including reservoirs of the Central Plains which often have both highly mixed (e.g. riverine zone) and thermally stratified (e.g. lacustrine zone) regions.
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