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Riffles 2 and 3 were located in an actively restored section of the river following dam removal i. Riffle 7 also existed before dam removal Full size image In total, we surveyed five riffles upstream of the previous dam location, one riffle below the previous dam location, and one riffle downstream of an existing lowhead dam in the same river, which served as a reference site Fig. Within each riffle, three quadrats were established at the upstream, middle, and downstream portions of the riffle to characterize representative microhabitats based on flow and substrate characteristics.
Fish, benthic macroinvertebrate, chemical water-quality, substrate, and streamflow surveys were conducted within each quadrat at six time intervals: late spring June , summer August , and late fall of ; early spring March , late spring June , and summer August of This is not a before-after study; in fact, riffles were non-existent or obscured in the impounded area before dam removal, making comparable benthic macroinvertebrate collections not feasible at riffles 2—6.
However, we collected macroinvertebrates downstream of the dam before removal riffle 7. Relative grain roughness for each quadrat was calculated using the ratio of streamflow depth divided by D84 i. Depth m was measured with a stadia rod as distance from the water surface to the substrate. Benthic macroinvertebrates and fish At each quadrat and time interval, a 0. To prevent fish from leaving the quadrat, a frame with a weighted net 4.
Pulling on upstream and downstream release cords enabled us to remotely set the frame net into final position. A time delay of 15 min between setting the frame and sampling the quadrat permitted a period without disturbance prior to sampling Bain et al. One electrofishing pass of s for each quadrat was conducted total of s per riffle. After collection and following enumeration and identification to species, fish were released.
Due to low fish numbers, only species richness, density, and number of darter species were calculated for fish assemblages. These metrics were calculated for each quadrat as well as for each riffle for each time period. The index represents the probability that two individuals randomly selected from a sample will belong to different species, or in this case, family. Linear mixed-effects models were used to test for differences in fish and benthic macroinvertebrate metrics among riffles over time, as well as for differences in streamflow velocity, mean water depth, and sediment-size distribution D16 and D50 and roughness.
Following Davis et al. For water-chemistry parameters, for which we only had one sample per reach per time period, one-way analysis of variance ANOVA was used to compare all reaches by time steps. Simple regression analysis was used to explore potential relationships between 1 roughness, D50, and changes in channel slope and benthic macroinvertebrate and fish metrics; 2 benthic macroinvertebrate density and fish density.
In NMS, distance matrices are rank-ordered and the solution determined iteratively by minimization of the stress criterion Kruskal The ANOSIM statistic R denotes the magnitude of the difference among groups; R equals 1 when groups differ completely and equals 0 when there is no difference detected among groups. Redundancy analysis RDA was used to identify potential differences in community composition of fish and benthic macroinvertebrate assemblages as a function of hydrogeomorphic and water-chemistry predictors among riffles i.
We used these metrics because of their importance to benthic macroinvertebrates and riffle fishes Kessler and Thorp ; Paul and Meyer and because they reasonably represented the variability observed in the broader set of streamflow and water-chemistry parameters surveyed as part of this study. Thus, RDA is useful for distinguishing the effects of dam removal on macroinvertebrate and fish assemblages in terms of drivers related to either hydrogeomorphology average stream flow velocity, D50 or chemical water quality DO, PO4.
Results Sediment-size distribution, water depth, and streamflow velocity varied by riffle and over time. The density and diversity of macroinvertebrates and fish were also different over time, largely as a function of season. Macroinvertebrate assemblage composition was different by time but not riffle, whereas fish assemblages were similar irrespective of time or riffle. Both hydrogeomorphic characteristics and chemical water-quality parameters emerged as potential drivers of macroinvertebrate and fish assemblages.
Riffle 7, downstream of the previous dam, generally exhibited the highest water temperatures as compared to the other riffles over time. Total P and PO4 were more consistent over time, but still exhibited differences among time periods note the high PO4 concentration at riffle 1 in August Across the study riffles and time periods, gravel ranged from Riffle 2 coarsened the most during the study period.
The substrate composition of our reference site—riffle 1 downstream riffle of an existing dam —remained fairly consistent across the study period. D16 and D50 varied among study riffles and through time Table 1 , Fig. D16 increased by 8. D50 was negatively associated with densities of both Chironomidae Fig.
Table 1 Linear mixed-effects models for fish and benthic macroinvertebrate response variables. Also included are D16 and D50, relative roughness, streamflow velocity, and mean water depth Full size table Fig. Riffle 1 is the reference riffle, riffle 2—6 are upstream of previous dam , and riffle 7 is downstream of the previous dam.
For visual clarity, error bars are not included. The lowest flows occurred in March , and the highest occurred in June although note missing values from November Water depth ranged from 0. Mean channel slope was 0. S2 a but negatively related to fish species richness Supplementary Material: Fig. S2 b and density Supplementary Material: Fig. Relative bed roughness was significantly different through time Table 1 , with the greatest decrease in roughness at riffle 4 and the largest increase in roughness at riffle 7.
Biotic assemblages Benthic macroinvertebrate density averaged For comparison, density was individuals 0. Macroinvertebrate density was also different over time Table 1 ; Fig. As a point of reference, macroinvertebrate abundance at riffle 1 as assessed by the Ohio EPA was variable between and , and the abundance in was within the previous range and on par with and Supplementary Material: Fig.
Riffle 1 is the reference riffle, riffles 2—6 are upstream of previous dam , and riffle 7 is downstream of the previous dam. S3 for details on data variability Full size image Fish density across all riffles at all time steps averaged 0. Fish density was also significantly different by time Fig. Fish density was not different among study riffles Table 1. Thirty insect families were represented in 12 orders, as well as class Oligochaeta and phylum Platyhelminthes.
Benthic macroinvertebrate richness was lowest in March across the year, but was comparable across the study riffles in June and June Fig. Across six of the seven sites except riffle 1 , there was a decrease in macroinvertebrate density from August to August Macroinvertebrate richness was not different among study riffles, but was significantly different through time Table 1. Macroinvertebrate evenness also varied significantly over time Supplementary Material: Table S1 , showing greatest evenness at riffle 4 and lowest at riffle 6 data not shown.
Fish species richness and darter species richness both ranged from 0 to 3 by study riffle. Richness of all fish species Fig. Rainbow and Banded Darters were found at all study sites except for riffle 7. Greenside Darters E. Bluebreast Darters E. For fish assemblages, there was no difference in composition by time Fig. The stress values were 0. The different shapes indicate the different riffles and the different colors indicate the different sampling time periods; only the most significant families are indicated.
Note that there are differences in the temporal representation of fish data in b i. For species and site scores, species were scaled proportionally to associated eigenvalues, while sites remained unscaled. Across all study riffles over time, RDA showed that benthic macroinvertebrate abundance had a weak positive association with D50 data not shown.
PO4 was at least weakly and positively related to benthic macroinvertebrate density in all but riffles 1 and 5 mostly in late spring and summer; see Supplementary Material: Fig. DO positively aligned with benthic macroinvertebrate family richness riffles 1, 2, and 6; see Supplementary Material: Fig. S5 a, b, f but not density but note negative relationship at riffle 5; Supplementary Material: Fig.
S5 e for certain time periods. Of the physicochemical variables, streamflow velocity emerged as the most influential physicochemical variable from the RDA, where it aligned with macroinvertebrate family richness or density at multiple riffles, but was inconsistent relative to the nature of the association Supplementary Material: Fig. Although we detected no differences in fish assemblages by time or riffle, fish and darter species richness were positively associated with D16, D50, and streamflow velocity at multiple riffles and time periods e.
Blue arrows indicate how environmental variables were ordinated. Where possible, riffles and time steps e. For locations where labels would overlap and be illegible, letters A, B, C, D, and E represent the respective riffles and time steps as listed in the legend Discussion Riffle development following dam removal Altered streamflow patterns characteristic of impounded rivers destroy riffles within the reservoir pool and impede the maintenance of riffles both further upstream as well as downstream.
After dam removal, new riffles can form in previously impounded reaches, although these bedforms may exhibit low habitat diversity compared to reference riffles Burroughs et al. We observed that five riffles had developed upstream of the previous dam in the former pool and its tail; riffles 2—6. Two of the upstream reaches were within an area of active channel restoration, likely leading to rapid development. Similar to other studies e.
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