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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.

Tennessee↗

Temperature and precipitation history of the Arctic

As the planet cooled from peak warmth in the early Cenozoic, extensive Northern Hemisphere ice sheets developed by 2.6 Ma ago, leading to changes in the circulation of both the atmosphere and oceans. From ∼2.6 to ∼1.0 Ma ago, ice sheets came and went about every 41 ka, in pace with cycles in the tilt of Earth’s axis, but for the past 700 ka, glacial cycles have been longer, lasting ∼100 ka, separated by brief, warm interglaciations, when sea level and ice volumes were close to present. The cause of the shift from 41 ka to 100 ka glacial cycles is still debated. During the penultimate interglaciation, ∼130 to ∼120 ka ago, solar energy in summer in the Arctic was greater than at any time subsequently. As a consequence, Arctic summers were ∼5 °C warmer than at present, and almost all glaciers melted completely except for the Greenland Ice Sheet, and even it was reduced in size substantially from its present extent. With the loss of land ice, sea level was about 5 m higher than present, with the extra melt coming from both Greenland and Antarctica as well as small glaciers. The Last Glacial Maximum (LGM) peaked ∼21 ka ago, when mean annual temperatures over parts of the Arctic were as much as 20 °C lower than at present. Ice recession was well underway 16 ka ago, and most of the Northern Hemisphere ice sheets had melted by 6 ka ago. Solar energy reached a summer maximum (9% higher than at present) ∼11 ka ago and has been decreasing since then, primarily in response to the precession of the equinoxes. The extra energy elevated early Holocene summer temperatures throughout the Arctic 1–3 °C above 20th century averages, enough to completely melt many small glaciers throughout the Arctic, although the Greenland Ice Sheet was only slightly smaller than at present. Early Holocene summer sea ice limits were substantially smaller than their 20th century average, and the flow of Atlantic water into the Arctic Ocean was substantially greater. As summer solar energy decreased in the second half of the Holocene, glaciers re-established or advanced, sea ice expanded, and the flow of warm Atlantic water into the Arctic Ocean diminished. Late Holocene cooling reached its nadir during the Little Ice Age (about 1250–1850 AD), when sun-blocking volcanic eruptions and perhaps other causes added to the orbital cooling, allowing most Arctic glaciers to reach their maximum Holocene extent. During the warming of the past century, glaciers have receded throughout the Arctic, terrestrial ecosystems have advanced northward, and perennial Arctic Ocean sea ice has diminished. Here we review the proxies that allow reconstruction of Quaternary climates and the feedbacks that amplify climate change across the Arctic. We provide an overview of the evolution of climate from the hot-house of the early Cenozoic through its transition to the ice-house of the Quaternary, with special emphasis on the anomalous warmth of the middle Pliocene, early Quaternary warm times, the Mid Pleistocene transition, warm interglaciations of marine isotope stages 11, 5e, and 1, the stage 3 interstadial, and the peak cold of the last glacial maximum.

Quaternary Science Reviews↗

Guadalupe Bass flow-ecology relationships; with emphasis on the impact of flow on recruitment

Guadalupe Bass Micropterus treculii is an economically and ecologically important black bass species endemic to the Edwards Plateau ecoregion and the lower portions of the Colorado River in central Texas. It is considered a fluvial specialist and as such, there are concerns that the increasing demands being placed upon the water resources of central Texas by growing human populations have the potential to negatively impact Guadalupe Bass populations. Therefore, this study assessed the relationship between Guadalupe Bass growth, feeding ecology, and streamflow. Sagittal otoliths were removed from Guadalupe Bass collected from throughout their range during 2015-2017 and used to estimate the age and back-calculate the growth trajectory of each individual. Additionally, young-of-year (YOY) Guadalupe Bass were collected every 10-14 days from two second-order streams, the North Llano River and South Llano River, in the Colorado Basin on the Edwards Plateau. Stomach contents of these individuals were identified and the effect of streamflow on the occurrence of the taxa comprising the stomach contents assessed. Guadalupe Bass growth was greater in the Colorado and Guadalupe River basins, independent of stream order, and tended to increase with increasing stream order within a basin. Growth was higher in higher stream orders and during years with stable and lower spring and summer monthly median flows, lower minimum and maximum flows, slower rise and fall rates, and higher baseflows. Growth was not influenced by years with higher monthly median flows in winter. These results would seem to contradict previous research, but more likely represent a fuller picture of how Guadalupe Bass respond to flow conditions. The disagreement between the current study and past studies seem to be attributable, at least in part, to the fact that previous studies were conducted during a period of extensive drought, while the current study was conducted during relatively wet conditions. Taken together with previous studies, the current study suggests that Guadalupe Bass growth is sensitive to flow conditions and is lower in years with flow conditions that fall outside a basin- and stream order-specific optimal range for the species. A total of 21 unique taxonomic groups were recovered from the stomachs of YOY Guadalupe Bass collected from the North Llano River and South Llano River. Aquatic insects, especially larval mayflies (Ephemeroptera), damselflies (Odanata: Zygoptera), and caddisflies (Trichoptera), were the most frequently encountered taxa. While there was no difference between the two rivers in stomach content composition, there was a strong longitudinal gradient in both systems with aquatic insects predominating at upstream sample sites and fishes being more common at downstream sites. Stream discharge during the 24 hours prior to collection did not have any influence on the probability of a taxa being found in Guadalupe Bass stomachs. The results of this study support efforts to manage Guadalupe Bass populations at a sub-watershed scale and suggests that populations occupying the same stream order within a basin are likely to have similar responses to annual flow conditions. In addition, these results indicate that the lower Colorado River population may inhabit a unique set of conditions that has supported the development of a trophy Guadalupe Bass fishery. Further, this study highlights the need to incorporate a sufficient range of annual flow conditions to ensure that the influence of stream flow on fish growth is adequately assessed. While interannual variation in growth rates seem to be capable of serving as a proxy for recruitment and year-class strength, long-term monitoring of recruitment paired with assessment of growth is necessary to further clarify the relationship between population density, flow regime, recruitment and growth and allow the construction of predictive models.

Texas↗

The geography and geology of Alaska; a summary of existing knowledge, with a section on climate, and a topographic map and description thereof

Alaska, the largest outlying possession of the United States, is that great land mass forming the northwestern extremity of the North American continent, whose western point is within 60 miles of the Asiatic coast (PI. II). About one-quarter of this area lies within the Arctic Circle, and from the standpoint of geographic position must be regarded as an arctic province; but the southern seaboard, exposed to the warm winds and waters of the Pacific, gives to the entire southern portion of the territory" a comparatively warm climate. It is not generally realized that the range of climate in Alaska is greater than that between Florida and Maine. At the southernmost point of the Pacific coast the mean annual temperature is not far from that of the city of Washington, the winters being warmer and characterized by less snowfall; the Yukon Valley on the other hand has a winter climate similar to that of northern Montana and Dakota; while in the extreme northern part of the territory the meteorologic conditions are invariably arctic. Though as yet only sparsely settled, Alaska's vast area and great resources make it one of the most important possessions of the United States and promise its rapid development. During the years 1890 to 1900 the population increased from 32,052* to 63,592. The mineral output, which in 1890 was valued at less than $800,000, exceeded $9,000,000 in 1904, and the fisheries show a corresponding growth. This rapid development has attracted public attention and led to urgent demand for explorations, surveys, and other investigations. So actively has this work been pushed, both by public and private enterprise, that exact knowledge of the geography, geology, and mineral resources of the interior has made greater strides within the last eight years than during the preceding thirty-one years since the acquisition of Alaska. The facts regarding the geography and geology, scattered as they are through the many books and reports of this period, are not always readily accessible, and the time seems ripe to present them in a summarized form. The topography of Alaska is varied and complex (see PI. I), and it is not easy to present briefly even the salient features. The limited number of pages here devoted to the subject precludes the possibility of detailed treatment, even if the facts were available. Much of the description has been taken from the results attained by other investigators, the writer being personally familiar with only a part of this large province. A list of the publications consulted is appended. The larger geographic features of Alaska are now fairly well known, though the detailed surveys which are demanded by the development of many localities have hardly been begun. Preliminary surveys have been completed of all but three 8 of the larger rivers. The most important mountain ranges have been at least outlined (fig. 3). Only three large areas remain almost entirely unmapped: One in southwestern Alaska, between Cook Inlet and the lower Kuskokwim, and the others in northern Alaska, embracing the Arctic watershed east and west of the Colville River. Nearly all the surveys of the interior, however, have been of a preliminary and exploratory character, and to meet the requirements of exact geography must be followed by more detailed mensuration. Though the coast line has been fairly well known for more than half a century, knowledge of the interior has been gained chiefly within the last two decades. This has not yet found its way into text-books and has too often been entirely ignored by cartographers. If facts are presented which may seem elementary, it is because even well-informed people have been known to harbor misconceptions in regard to the orographic features, climate, and general character of Alaska. Those who read of the perils and privations of winter travel and explorations are apt to picture a region of ice and snow; others, again, who have personal knowledge of the tourist route of southeastern Alaska, regard the whole district as one of rugged mountains and glaciers. In point of fact, glaciers are now nearly limited to the ranges bordering the Pacific and to the two slopes of the Alaska range; and even during the greatest development of glaciers but a small portion of Alaska was under ice (see map, PI. XXII). As a treatise on geography would hardly be complete without some discussion of the climate, meteorologic data have been compiled by Mr. Cleveland Abbe, jr., but the discussion of this does not pretend to be more than a cursory treatment of the subject. The scope of the paper seems to require also a brief summary of the development of geographic knowledge of Alaska. This subject, with its many ramifications, is of fascinating interest and offers a magnificent field for the trained historian. If the accompanying sketch of discovery and exploration awakens any measure of popular interest the writer will feel amply rewarded for having attacked a theme which hardly falls within the scope of his investigations. When this compilation was begun it was intended to be chiefly a description of the topography of Alaska, as illustrated by the accompanying map (PI. XXXIV, in pocket), which was compiled under the direction of the late R. U. Goode. In the course of the work there accumulated much geologic as well as geographic material which seemed worthy of inclusion in the report. As no comprehensive statement of the geology of Alaska has been made since the modern epoch of investigation was begun, an attempt will be made to give a summary of all results achieved. Since the writer has obtained much of his knowledge of the facts from the work of others, he disclaims any pretense of making an entirely original contribution to geologic science. He feels, however, that a personal familiarity with a considerable part of the province, gained during seven consecutive seasons of field work, will justify Mm in presenting conclusions which may in some cases be at variance with those in the reports on which he must draw for his facts. Throughout this report attempt will be made to credit borrowed material to the source from which it is drawn. Where such matter has been obtained entirely from published reports there is no difficulty in so doing; but as regards investigators of the Geological Survey, with whom the writer has collaborated both in field and in office, the case is somewhat different, for it is not always possible to know whether this or that theory originated with the writer or with one of his colleagues. It will, then, perhaps suffice to state that this report could not have been prepared without the explorations and researches of the geologists, F. C. Schrader, Walter C. Mendenhall, Arthur J. Collier, J. E. Spurr, and Arthur C. Spencer; and the surveys of the topographers, T. G. Gerdine, D. C. Witherspoon, D. L. Reaburn, W. J. Peters, and E. C. Barnard. Each of these men, in the course of from two to six years of field work, has made important contributions to the knowledge of the geography and geology of Alaska, and not all of these results have yet been put in print. In the last season (1903) L. M. Prindle, C. W. Wright, Arthur Hollick, G. C. Martin, F. L. Hess, and Fred H. Moffit have carried on geologic work in Alaska, and the writer has made use of their work now in course of publication. He has also been fortunate in having access to the manuscript reports of Walter C. Mendenhall and F. C. Schrader on the Copper River basin, to which references will be made. The matter here presented should be credited in a measure to all of these investigators, but for many of the theories advanced the writer alone is responsible. As this manuscript goes to press there has been opportunity to incorporate some of the results of the field work of 1904. As far as possible these have been embodied in the text, but in some instances it has been found advisable to add them only as footnotes. During the past summer F. E. and C. W. Wright extended the geologic reconnaissance in southeastern Alaska. In southwestern Alaska G. C. Martin and T. W. Stanton have determined the general Mesozoic section, while F. H. Moffit has made a reconnaissance of the northern part of the Kenai Peninsula. A. J. Collier has mapped the geology of the Cape Lisburne region, and L. M. Prindle and F. L. Hess have made contributions to the knowledge of the metamorphic terranes of the Yukon-Tanana district. It is the writer's purpose to describe in nontechnical language the larger geographic features and discuss their relation as far as the data available will permit. In the treatment of the geology, however, less effort will be made to make the matter acceptable to the lay reader. It is hoped, however, that a brief summary of the salient features of the geologic history' may be not without interest to the general public. If this paper serves in some measure to dispel the popular fallacies regarding Alaska and to disseminate more accurate knowledge of its geographic and geologic features, the purpose of its publication will be accomplished.

Alaska↗

The Missoula and Bonneville floods—A review of ice-age megafloods in the Columbia River basin

The Channeled Scabland of eastern Washington State, USA, brought megafloods to the scientific forefront. A 30,000-km2 landscape of coulees and cataracts carved into the region’s loess-covered basalt attests to overwhelming volumes of energetic water. The scarred landscape, garnished by huge boulder bars and far-travelled ice-rafted erratics, spurred J Harlen Bretz’s vigorously disputed flood hypothesis in the 1920s. First known as the Spokane flood, it was rebranded the Missoula flood once understood that the water came from glacial Lake Missoula, formed when the Purcell Trench lobe of the last-glacial Cordilleran ice sheet dammed the Clark Fork valley in northwestern Idaho with ice a kilometer thick. Bretz’s flood evidence in the once-remote Channeled Scabland, widely seen and elaborated by the 1950s, eventually swayed consensus for cataclysmic flooding. Missoula flood questions then turned to some that continue today: how many? when? how big? what routes? what processes? The Missoula floods passed through eastern Washington by a multitude of valleys, coulees and scabland tracts, some contemporaneously, some sequentially. Which routings and their timing depended on the positions of various lobes of the multi-pronged Cordilleran ice sheet and the erosional development of the channels themselves. The first floods mostly followed the big bend of Columbia valley looping through north-central Washington. But the south-advancing Okanogan ice lobe soon blocked that path, forming long-lasting glacial Lake Columbia in the impounded Columbia valley. Missoula floods into this lake were diverted south out of the Columbia valley and into eastern Washington coulees and scabland tracts. At least four floods entered Moses Coulee, but then as the Okanogan lobe advanced over and blocked the head of that coulee, more eastern paths took the water, including Grand Coulee and the Telford-Crab-Creek and Cheney-Palouse scabland tracts. Flood routing also depended on the erosion of the coulees. At some point, headward erosion of upper Grand Coulee lowered the divide saddle between the west-running Columbia valley and the deep and wide Grand Coulee heading southwest. Still uncertain is when this happened and the consequences with respect to the stage and extent of glacial Lake Columbia and to flood access to the other, higher, flood routes. Downstream, all flood routes converged onto Pasco Basin, flowed through Wallula Gap and the Columbia River Gorge into the Pacific Ocean, following submarine canyons and depositing sediment layers on abyssal plains. Stratigraphic studies indicate dozens—likely more than a hundred—separate Missoula floods during the last glacial period. Over the length of the flood route, backwater areas and depositional basins preserve multiple flood beds, many of which are separated by signs of time, including volcanic ash layers and soil development in subaerial environments; and varve-like beds and pelagic mud layers in lacustrine and marine settings. Evidence also comes from the glacial Lake Missoula basin, where stratigraphy indicates dozens of filling and emptying cycles. Varve counts in conjunction of radiocarbon dating and paleomagnetic secular variation show the repeated filling-and-release cycles of glacial Lake Missoula had intervals possibly as long as 100 years early in the lake’s history but diminished to just one or two years for the last few floods. This behavior accords with jökulhlaup-style floods released by subglacial drainage from a self-dumping ice-dammed lake. But not yet clear is whether such a mechanism applies to all the floods or if some emptied more cataclysmically as hypothesized by some. Radiocarbon dating of sparse organic materials remains key to defining flood chronology but has been lately bolstered by analyses of terrestrial cosmogenic nuclides and optically stimulated luminescence. Varve counts and paleomagnetic secular variation studies help to define durations and intervals represented by sequences of flood beds. The ~16 ka Mount St. Helens Set S tephra is commonly interbedded within flood deposits, enabling correlation of deposits among sites. Tephra from the 13.7–13.4 ka eruption of Glacier Peak overlies all glacial Lake Missoula and Missoula flood deposits, defining an end time. Overall conclusions are that glacial Lake Missoula was extant and producing floods for at least 3–4 ky during 20–14 ka. At least ~75 floods preceded Mount St Helens Set S, followed by 30 or more after the tephra fall. Most floods entered glacial Lake Columbia, impounded by the Okanogan lobe, for 2–5 ky between about 18.5 and 15 ka. Glacial Lake Columbia outlived Lake Missoula by >200–400 yr but may have been born later since at least one flood came down the Columbia valley before the Okanogan ice lobe blocked the Columbia valley at 18.5–18 ka. The maximum extent of the Okanogan and Purcell Trench lobes, many Missoula floods, substantial erosion of upper Grand Coulee, and the widespread tephra falls from Mount St. Helens eruptions all happened about 17–15 ka. People, in the area since 16.6–15.3 ka, almost certainly witnessed the last of the Missoula floods and later large floods from other ice-dammed lakes in the Columbia River basin. Quantitative flow analyses give peak discharge estimates and support understanding of erosional and depositional processes. The first flow assessments were simple cross-section calculations but recent assessments employ two-dimensional hydrodynamic models. The general finding is that emplacement of the maximum stage evidence requires about 20 million m3/s near the Lake Missoula outlet and about 5–15 million m3/s through Wallula Gap and downstream in the Columbia River Gorge. These hydraulic analyses raise still-unresolved questions regarding canyon erosion and possible additional water sources. The large Pleistocene Bonneville flood entered the Columbia River system from the southeast from pluvial Lake Bonneville, the Pleistocene predecessor to Great Salt Lake in the eastern Great Basin. During the last glacial, the lake basin filled, covering >50,000 km2 with 10,400 km3 of water before reaching its maximum possible stage governed by Red Rock Pass, the lowest divide separating the basin from the Snake River basin to the north. The overtopping lake rapidly incised 108–125 m into the Red Rock Pass outlet, spilling half of its total lake volume. G.K. Gilbert described the essential sequence in the 1870s, but the flood was mostly forgotten until the late 1950s when Harold Malde linked the spectacular scabland topography and bouldery “melon gravel” on the Snake River Plain to the Lake Bonneville overflow. The Bonneville flood appears to have been a singular event at about 18 ka. No evidence of multiple or pre-last-glacial spillovers has yet been found. Its total volume was about twice that of a maximum Lake Missoula flood yet its peak discharge was ~1 million m3/s, less than a tenth of the largest Missoula floods. Its comparatively simple flow path and much steadier flow make the Bonneville flood ideal for new studies of erosional and depositional processes. At least two floods seem to have passed down the Columbia valley after the last of the Missoula floods, including a large flood about ~14 ka likely from cataclysmic demise of the thinning Okanogan ice lobe dam impounding glacial Lake Columbia. Floods from earlier glacial ages left scant yet clear evidence in the Channeled Scabland and Columbia valley. But their source, timing, and magnitudes are little understood. Some deposits are paleomagnetically reversed, thus older than ~800 ka. Last-glacial floods and perhaps older ones affected the Snake River Plain, some likely sourced in lakes dammed by alpine glaciers in central Idaho.

Idaho, Oregon, Washington↗