Small creatures, big challenge: A review of current knowledge and assessment of research needs to inform water withdrawal management for the Duck River's freshwater mussel fauna

The Duck River is a remarkable national resource, boasting outstanding biodiversity (Ahlstedt et al. 2017). It contains 151 fish species, over 60 mussel species, and 22 snail species, many of which are federally listed (Etnier and Starnes 1993; Johnson et al. 2013; Ahlstedt et al. 2017; Womble and Rosenberger 2025a). The Duck River watershed is at a pivotal time in its management and ecological history due to significant growth in human population and economic activity. The consequent increase in surface water demand presents challenges for sustainable water resource management. The Tennessee Department of Environment and Conservation (TDEC) has permitted or proposed to permit eight water utilities to withdraw over 77 million gallons per day (mgd) from the Duck River, representing > 40% increase from previously permitted amounts (USFWS 2025). On November 20, 2024, Tennessee Governor Bill Lee signed Executive Order No 108 to ensure long-term stability of the Duck River watershed through balancing economic growth, water resource management, and environmental conservation. Some primary components of this plan are the creation of a Comprehensive Regional Drought Management Plan, the development of a Habitat Conservation Plan, and the establishment of the Duck River Watershed Planning Partnership (DRWPP). Although the planning partnership is not focused on a sole aspect of the Duck River, but the watershed as a whole, the mussel assemblages of the Duck River have been a particular focus due to their importance in providing ecosystem services, sensitivity to water quality, conservation status, and sensitivity to periodic drought conditions in this system. 

With multiple state and federal agencies involved in managing the Duck River’s unique water resources, a focused review on how future water management could affect the freshwater mussel communities in the Duck River will contribute to this process. 

Our objectives of this review are to:

1. Review the effects of changing flow regimes on mussels, with emphasis on the effect of droughts/low flows, highlighting key processes/factors affecting mussels.

2. Highlight critical data gaps that could allow future, evidence-based water management decisions in this system.

3. Propose research topics to address critical data gaps and enable evidence-based water management decisions that consider the needs of freshwater mussels. 

Freshwater mussels are a critical component of riverine ecosystems, providing ecosystem services including water filtration, nutrient cycling, and habitat provision (Vaughn et al. 2004; Haag 2012) that benefit human communities. Environmental flows, also referred to as ecological or e-flows, refer to the quantity, timing, and quality of water flows necessary to maintain the ecological health of rivers, wetlands, and other aquatic ecosystems (Acreman and Dunbar 2004; Acreman 2016). For freshwater mussels, environmental flows include maintaining a flow regime and associated water quality regimes that are favorable for both juveniles and adults while also seasonally supporting host fish and the survival of settling glochidia (i.e., larval mussels) in suitable environments (Gates et al. 2015). Freshwater mussels are adapted to the natural flow regime of their native streams, including seasonal variability in flows. As sedentary filter feeders, mussels require adequate water flow to provide oxygen and food (Allen and Vaughn 2009); however, they also require periods of low flow to increase access to fish hosts and to ensure juveniles are not displaced (Gates et al 2015). Mussels are sensitive to water quality conditions, including pollutants, sedimentation, and artificial alterations of temperature and pH (Strayer and Malcom 2012). Mussel community characteristics, including presence, abundance, and species richness, generally increase with stream size (e.g., stream order); however, responses to changes in flow conditions are often species-specific, and the magnitude and timing of extreme flow conditions have more predictive power for mussel community response than general changes in flow (Lopez and Vaughn 2021). Complex hydraulic variables, such as relative shear stress and Reynolds number, can be strong predictors of mussel community responses because they combine simple hydraulic variables (e.g., velocity, depth) with stream channel characteristics, such as substrate composition and dynamics. 

Droughts directly and indirectly affect mussels by influencing their physiology, behavior, reproduction, and ecosystem roles. Mass mussel die-offs during droughts and low water conditions are widely documented (Vaughn et al. 2015; Sousa et al. 2018; Dubose et al. 2019). Droughts can also lead to a shift in mussel community composition, with drought-tolerant species becoming more dominant while more sensitive species experience declines (Galbraith et al, 2010; Lopez et al. 2022; Vaughn and Atkinson 2025). Droughts affect mussels through direct mortality from stranding as well as physiological stress from altered water temperature and quality, reduced habitat and food availability and quality, increased predation and competition, disruption of reproductive cycles, and increased susceptibility to disease and pathogens. Although the effects of droughts, water withdrawals, and low flow conditions on mussels are an increasing area of study, there is a general paucity of data-driven consensus regarding optimal mitigation approaches and strategies (Cushway et al. 2024). Long-term management decisions regarding water withdrawals, particularly during extended or multi-year drought conditions, may consider factors beyond direct mortality such as the sublethal effects of drought and low flow conditions on freshwater mussels for a comprehensive understanding of the effects of prolonged low flows on freshwater mussels and aquatic communities.