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Department of Biological Sciences


Species invasions are changing aquatic ecosystems worldwide. Submerged aquatic macrophytes control lake ecosystem processes through their direct and indirect interactions with other primary producers, but how these interactions may be altered by macrophyte species invasions in temperate lakes is poorly understood. We addressed whether invasive watermilfoil (IWM) altered standing crops and gross primary production (GPP) of other littoral primary producers (macrophytes, phytoplankton, attached algae, and periphyton) in littoral zones of six Michigan lakes through a paired-plot comparison study of sites with IWM (standardized abundance 7–56%) compared to those with little or no IWM (standardized abundance 0–2%). We found that primary producer standing crops and the GPP of epiphytes, phytoplankton, and benthic periphyton were variable among lakes and not significantly different between paired study plots. Macrophyte standing crops predicted rates of benthic periphyton GPP, and standing crops of all other primary producers across all study plots. Overall, our results suggest that the effects of IWM on other primary producers in littoral zones may be lake-specific, and are likely dependent on the density of IWM, or whether it is functionally similar to other native species that it replaces or co-exists with. Moreover, in lakes where IWM is established but does not dominate macrophyte assemblages, the effects on littoral zone productivity may be minimal. Instead, overall macrophyte biomass is the primary factor controlling the rates of production and biomass of the other littoral zone primary producers, as has long been understood and observed in lake ecosystems.

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© 2020 by the authors. Licensee MDPI, Basel, Switzerland. This article is an open access article distributed under the terms and conditions of the Creative Commons Attribution (CC BY) license ( Publisher’s version of record:

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Creative Commons Attribution 4.0 International License
This work is licensed under a Creative Commons Attribution 4.0 International License.


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