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Ecosystem bioelement variability is associated with freshwater animal aggregations at the aquatic-terrestrial interface

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The impacts of animals on the biogeochemical cycles of major bioelements like C, N, and P are well-studied across ecosystem types. However, more than 20 elements are necessary for life. The feedbacks between animals and the biogeochemical cycles of the other bioelements are an emerging research priority. We explored how much freshwater mussels (Bivalvia: Unionoida) were related to variability in ecosystem pools of 10 bioelements (Ca, Cu, Fe, K, Mn, Na, Mg, P, S and Zn) in streams containing a natural mussel density gradient in the US Interior Highlands. We studied the concentrations of these bioelements across the aquatic-terrestrial interface—in the porewater of riverine gravel bars, and the emergent macrophyte Justicia americana. Higher mussel density was associated with increased calcium in gravel bars and macrophytes. Mussel density also correlated with variability in iron and other redox-sensitive trace elements in gravel bars and macrophytes, although this relationship was mediated by sediment grain size. We found that two explanations for the patterns we observed are worthy of further research: (1) increased calcium availability in gravel bars near denser mussel aggregations may be a product of the buildup and dissolution of shells in the gravel bar, and (2) mussels may alter redox conditions, and thus elemental availability in gravel bars with fine sediments, either behaviorally or through physical structure provided by shell material. A better understanding of the physical and biogeochemical impacts of animals on a wide range of elemental cycles is thus necessary to conserve the societal value of freshwater ecosystems.

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The datasets used and analyzed during the current study are uploaded to Open Science Framework and can be found here (


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We thank A. Franzen, E. Higgins, the members of the Vaughn Lab, A. Cooper, M. Spikes, and P. Jeyasingh for assistance. We thank L. Souza, M. Kaspari, B. Hall, and an anonymous reviewer for comments that improved the manuscript. J. Hartwell designed Fig. 1. We also thank the landowners that allowed us access to our study rivers through their property. This paper is part of a dissertation at the University of Oklahoma and a contribution to the program of the Oklahoma Biological Survey.


Funding for this work was provided by the Oklahoma Department of Wildlife Conservation, the University of Oklahoma, Department of Biology, the Oklahoma Biological Survey, Western Ag Innovations, and the National Science Foundation (DEB 1457542).

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Authors and Affiliations



JWL and CCV designed the field studies. RNH conducted and advised on ICP-OES analyses. TBP assisted with the PRS® probe study and consulted on data analysis and interpretation. JWL wrote the initial manuscript and revised with input from all authors.

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Correspondence to Jonathan W. Lopez.

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The authors declare that they have no conflict of interest.

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Communicated by Robert O. Hall.

We used a multi-element approach to examine relationships between animal aggregations and ecosystems. This approach has potential to generate hypotheses regarding animals' biogeochemical impacts.

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Lopez, J.W., Hartnett, R.N., Parr, T.B. et al. Ecosystem bioelement variability is associated with freshwater animal aggregations at the aquatic-terrestrial interface. Oecologia 202, 795–806 (2023).

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