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Factors affecting the occurrence of saugers in small, high-elevation rivers near the western edge of the species' natural distribution

Factors affecting the occurrence of saugers Sander canadensis were studied throughout the Wind River basin, a high-elevation watershed (> 1,440 m above mean sea level) on the western periphery of the species' natural distribution in central Wyoming. Adult saugers appeared to have a contiguous distribution over 170 km of streams among four rivers in the watershed. The upstream boundaries of sauger distribution were influenced by summer water temperatures and channel slopes in two rivers and by water diversion dams that created barriers to upstream movement in the other two rivers. Models that included summer water temperature, maximum water depth, habitat type (pool or run), dominant substrate, and alkalinity accounted for the variation in sauger occurrence across the watershed within the areas of sauger distribution. Water temperature was the most important basin-scale habitat feature associated with sauger occurrence, and maximum depth was the most important site-specific habitat feature. Saugers were found in a larger proportion of pools than runs in all segments of the watershed and occurred almost exclusively in pools in upstream segments of the watershed. Suitable summer water temperatures and deep, low-velocity habitat were available to support saugers over a large portion of the Wind River watershed. Future management of saugers in the Wind River watershed, as well as in other small river systems within the species' native range, should involve (1) preserving natural fluvial processes to maintain the summer water temperatures and physical habitat features needed by saugers and (2) assuring that barriers to movement do not reduce upstream boundaries of populations.

Transactions of the American Fisheries Society

Application of neural networks to prediction of fish diversity and salmonid production in the Lake Ontario basin

Diversity and fish productivity are important measures of the health and status of aquatic systems. Being able to predict the values of these indices as a function of environmental variables would be valuable to management. Diversity and productivity have been related to environmental conditions by multiple linear regression and discriminant analysis, but such methods have several shortcomings. In an effort to predict fish species diversity and estimate salmonid production for streams in the eastern basin of Lake Ontario, I constructed neural networks and trained them on a data set containing abiotic information and either fish diversity or juvenile salmonid abundance. Twenty percent of the original data were retained as a test data set and used in the training. The ability to extend these neural networks to conditions throughout the streams was tested with data not involved in the network training. The resulting neural networks were able to predict the number of salmonids with more than 84% accuracy and diversity with more than 73% accuracy, which was far superior to the performance of multiple regression. The networks also identified the environmental variables with the greatest predictive power, namely, those describing water movement, stream size, and water chemistry. Thirteen input variables were used to predict diversity and 17 to predict salmonid abundance.

Transactions of the American Fisheries Society

Comparison of prehatch C-start responses in rainbow trout and lake trout embryos by means of a tactile stimulus test

The C-start in teleost fishes, a type of startle response, mediates the ability to respond to abrupt, unexpected stimuli and is characterized by a short-latency, C-type fast start acceleration. In prehatch fish embryos, the C-start appears necessary for mechanical breakdown of the egg chorion and successful hatching by way of increased embryo movement and distribution of the hatching enzymes. In later stages, the C-start plays an important role in predator avoidance. Using tactile stimulation, we evaluated the C-start response in rainbow trout Oncorhynchus mykiss at 170 degree-days, when 6.6% of embryos exhibited C-starts, and lake trout Salvelinus namaycush embryos at 320 degree-days, when 23% of embryos exhibited C-starts. Triplicate groups of embryos were later tested at three developmental stages: early (220 and 360 degree-days for rainbow trout and lake trout, respectively), middle (260 and 480 degree-days, respectively), and late (320 and 560 degree-days, respectively). The proportion of trout embryos exhibiting C-start increased through time, such that 100% had responded by the late stage, just prior to hatching. C-starts could be obtained by repeated stimulation, and the relative activity of the embryos (based on the number of flexures per stimulus) also increased over time. Rainbow trout and lake trout showed very similar C-start responses at parallel developmental stages, and these patterns of response were similar to those reported in other fish species.

Transactions of the American Fisheries Society

Development of a multimetric index for assessing the biological condition of the Ohio River

The use of fish communities to assess environmental quality is common for streams, but a standard methodology for large rivers is as yet largely undeveloped. We developed an index to assess the condition of fish assemblages along 1,580 km of the Ohio River. Representative samples of fish assemblages were collected from 709 Ohio River reaches, including 318 "least-impacted" sites, from 1991 to 2001 by means of standardized nighttime boat-electrofishing techniques. We evaluated 55 candidate metrics based on attributes of fish assemblage structure and function to derive a multimetric index of river health. We examined the spatial (by river kilometer) and temporal variability of these metrics and assessed their responsiveness to anthropogenic disturbances, namely, effluents, turbidity, and highly embedded substrates. The resulting Ohio River Fish Index (ORFIn) comprises 13 metrics selected because they responded predictably to measures of human disturbance or reflected desirable features of the Ohio River. We retained two metrics (the number of intolerant species and the number of sucker species [family Catostomidae]) from Karr's original index of biotic integrity. Six metrics were modified from indices developed for the upper Ohio River (the number of native species; number of great-river species; number of centrarchid species; the number of deformities, eroded fins and barbels, lesions, and tumors; percent individuals as simple lithophils; and percent individuals as tolerant species). We also incorporated three trophic metrics (the percent of individuals as detritivores, invertivores, and piscivores), one metric based on catch per unit effort, and one metric based on the percent of individuals as nonindigenous fish species. The ORFIn declined significantly where anthropogenic effects on substrate and water quality were prevalent and was significantly lower in the first 500 m below point source discharges than at least-impacted sites nearby. Although additional research on the temporal stability of the metrics and index will likely enhance the reliability of the ORFIn, its incorporation into Ohio River assessments still represents an improvement over current physicochemical protocols.

Transactions of the American Fisheries Society

Variable migratory patterns of different adult rainbow trout life history types in a southwest Alaska watershed

Radiotelemetry was used to document population structure in adult rainbow trout Oncorhynchus mykiss from the Alagnak River, southwest Alaska. Rainbow trout (N = 134) longer than 440 mm were implanted with radio transmitters and tracked for varying periods from July 1997 to April 1999. Fifty-eight radio-tagged fish were tracked for sufficient duration (at least 11 months) to allow description of seasonal migratory patterns. Unique seasonal movements of fish suggested discrete, within-basin population structure. Telemetry data documented the existence of multiple migratory and nonmigratory groups of rainbow trout, indicating unique life history patterns. The observed groups consisted of what we defined as a lake-resident ecotype, a lake-river ecotype, and a riverine ecotype; the riverive ecotype demonstrated both highly migratory and nonmigratory movement behavior. Considerable variation in movement patterns was found within both the lake-river group and the river migratory group. Radio-tagged trout did not migrate between the two Alagnak watershed lakes in either year of the study, suggesting lake fidelity in the population structure. Alagnak River rainbow trout may have evolved the observed seasonal movement patterns to optimize winter thermal refugia and summer food availability of salmon eggs and carcasses.

Alaska

Immobilization thresholds of electrofishing relative to fish size

Fish size and electrical waveforms have frequently been associated with variation in electrofishing effectiveness. Under controlled laboratory conditions, we measured the electrical power required by five electrical waveforms to immobilize eight fish species of diverse sizes and shapes. Fish size was indexed by total body length, surface area, volume, and weight; shape was indexed by the ratio of body length to body depth. Our objectives were to identify immobilization thresholds, elucidate the descriptors of fish size that were best associated with those immobilization thresholds, and determine whether the vulnerability of a species relative to other species remained constant across electrical treatments. The results confirmed that fish size is a key variable controlling the immobilization threshold and further suggested that the size descriptor best related to immobilization is fish volume. The peak power needed to immobilize fish decreased rapidly with increasing fish volume in small fish but decreased slowly for fish larger than 75-100 cm 3. Furthermore, when we controlled for size and shape, different waveforms did not favor particular species, possibly because of the overwhelming effect of body size. Many of the immobilization inconsistencies previously attributed to species might simply represent the effect of disparities in body size.

Transactions of the American Fisheries Society

Effects of flow regulation on shallow-water habitat dynamics and floodplain connectivity

Our study examined the effects of flow regulation on the spatiotemporal availability of shallow habitat patches with slow current velocity (SSCV patches) and floodplain inundation in the unregulated Yellowstone River and the regulated Missouri River in Montana and North Dakota. We mapped representative sites and used hydraulic models and hydrograph data to describe the frequency and extent of floodplain inundation and the availability of SSCV habitat over time during different water years. In the Yellowstone River the distribution, location, and size of SSCV patches varied but followed an annual pattern that was tied to the snowmelt runoff hydrograph. There was less variation in patch distribution in the Missouri River, and the pattern of habitat availability was influenced by flow regulation. Regulated flows and their effects on channel morphology and patterns of vegetation establishment resulted in 3.0–3.5 times less area of inundated woody vegetation during normal and dry years in the Missouri River compared with the Yellowstone River. The differences we observed in SSCV patch dynamics between rivers may have implications for fish populations and community structure through affecting the survival of early life stages. At a larger scale, the smaller area of vegetation inundated in the Missouri River suggests that nutrient cycling and the ecological benefits associated with a moving littoral zone are reduced by the altered flow and sediment regime in that river. Accurate assessments of the effects of flow alteration and successful efforts to restore riverine ecosystems will require consideration of physical and biotic processes that operate at multiple spatial and temporal scales.

Transactions of the American Fisheries Society

Genetic assessment of strain-specific sources of lake trout recruitment in the Great Lakes

Populations of wild lake trout Salvelinus namaycush have been extirpated from nearly all their historical habitats across the Great Lakes. Efforts to restore self-sustaining lake trout populations in U.S. waters have emphasized the stocking of coded-wire-tagged juveniles from six hatchery strains (Seneca Lake, Lewis Lake, Green Lake, Apostle Islands, Isle Royale, and Marquette) into vacant habitats. Strain-specific stocking success has historically been based on estimates of the survival and catch rates of coded-wire-tagged adults returning to spawning sites. However, traditional marking methods and estimates of relative strain abundance provide no means of assessing strain fitness (i.e., the realized contributions to natural recruitment) except by assuming that young-of-the-year production is proportional to adult spawner abundance. We used microsatellite genetic data collected from six hatchery strains with likelihood-based individual assignment tests (IA) and mixed-stock analysis (MSA) to identify the strain composition of young of the year recruited each year. We show that strain classifications based on IA and MSA were concordant and that the accuracy of both methods varied based on strain composition. Analyses of young-of-the-year lake trout samples from Little Traverse Bay (Lake Michigan) and Six Fathom Bank (Lake Huron) revealed that strain contributions differed significantly from estimates of the strain composition of adults returning to spawning reefs. The Seneca Lake strain contributed the majority of juveniles produced on Six Fathom Bank and more young of the year than expected within Little Traverse Bay. Microsatellite markers provided a method for accurately classifying the lake trout hatchery strains used for restoration efforts in the Great Lakes and for assessment of strain-specific reproductive success.

Transactions of the American Fisheries Society

Test of a Power Transfer Model for Standardized Electrofishing

Standardization of electrofishing in waters with differing conductivities is critical when monitoring temporal and spatial differences in fish assemblages. We tested a model that can help improve the consistency of electrofishing by allowing control over the amount of power that is transferred to the fish. The primary objective was to verify, under controlled laboratory conditions, whether the model adequately described fish immobilization responses elicited with various electrical settings over a range of water conductivities. We found that the model accurately described empirical observations over conductivities ranging from 12 to 1,030 ??S/cm for DC and various pulsed-DC settings. Because the model requires knowledge of a fish's effective conductivity, an attribute that is likely to vary according to species, size, temperature, and other variables, a second objective was to gather available estimates of the effective conductivity of fish to examine the magnitude of variation and to assess whether in practical applications a standard effective conductivity value for fish may be assumed. We found that applying a standard fish effective conductivity of 115 ??S/cm introduced relatively little error into the estimation of the peak power density required to immobilize fish with electrofishing. However, this standard was derived from few estimates of fish effective conductivity and a limited number of species; more estimates are needed to validate our working standard.

Transactions of the American Fisheries Society

Growth and condition of alewives in Lake Michigan, 1984-2001

Diets of salmonines in Lake Michigan have been dominated by alewives Alosa pseudoharengus since the 1960s, and information on alewife population dynamics is critical to the management of salmonine fisheries. We monitored alewife size at age and condition ( K ) at several different locations in Lake Michigan during fall 1984–2001. Alewives were aged by enumerating annuli on otoliths. The results indicated that alewife length at age did not trend upward or downward between 1984 and the late 1990s but decreased from the late 1990s to 2001. Alewife weight at age was relatively constant between 1984 and the mid-1990s but decreased from the mid-1990s to 2001. Mean condition for a given alewife age was, on average, 13.7% higher during 1984–1994 than during 1995–2001. This decline in alewife condition was not a density-dependent response by the alewife population because alewife abundance trended neither upward nor downward during 1984–2001. The decline in alewife condition was possibly due to the lakewide decrease in the abundance of Diporeia spp. during the 1990s. Apparently, the availability of the large-bodied invertebrates Diporeia and Mysis spp. was an important regulator of adult alewife growth because alewives attained a substantially larger size in Lake Michigan, where these invertebrates were relatively important constituents of the adult alewife diet, than in Lake Ontario, where these invertebrates were not readily eaten by adult alewives. For age-2 or older females, mean length was 2–9 mm greater than for males. Alewife size at age and condition were slightly higher on the eastern side of Lake Michigan than on the western side.

Transactions of the American Fisheries Society

Nonlinear response of trout abundance to summer stream temperatures across a thermally diverse montane landscape

Stream temperature is a fundamental physical factor that affects the distribution and abundance of salmonids, but empirical inconsistencies exist regarding the nature of this relationship in wild populations. We sampled trout populations composed primarily of cutthroat trout Oncorhynchus clarki but also including brown trout Salmo trutta and brook trout Salvelinus fontinalis at 102 reaches on 24 first- to fourth-order streams across a thermally diverse montane landscape. Curves fit to scatterplots of density and biomass versus mean July-August stream temperatures suggested nonlinear, dome-shaped responses. Peaks occurred near mean stream temperatures of 12??C; x-intercepts were near 3??C and 21??C. We conclude that inconsistencies in previously reported temperature-abundance relationships for wild trout populations may have resulted from sampling only a subset of the thermal environments occupied by a species. Researchers analyzing this relationship should be cognizant of the range of temperatures studied and the expected form of the relationship over that range.

Transactions of the American Fisheries Society

Effect of stock size, climate, predation, and trophic status on recruitment of alewives in Lake Ontario, 1978-2000

The population of alewives Alosa pseudoharengus in Lake Ontario is of great concern to fishery managers because alewives are the principal prey of introduced salmonines and because alewives negatively influence many endemic fishes. We used spring bottom trawl catches of alewives to investigate the roles of stock size, climate, predation, and lake trophic status on recruitment of alewives to age 2 in Lake Ontario during 1978–2000. Climate was indexed from the temperature of water entering a south-shore municipal treatment plant, lake trophic status was indexed by the mean concentration of total phosphorus (TP) in surface water in spring, and predation was indexed by the product of the number of salmonines stocked and relative, first-year survival of Chinook salmon Oncorhynchus tshawytscha . A Ricker-type parent–progeny model suggested that peak production of age-1 alewives could occur over a broad range of spawning stock sizes, and the fit of the model was improved most by the addition of terms for spring water temperature and winter duration. With the addition of the two climate terms, the Ricker model indicated that when water was relatively warm in spring and the winter was relatively short, peak potential production of young was nine times higher than when water temperature and winters were average, and 73 times higher than when water was cold in spring and winters were long. Relative survival from age 1 to recruitment at age 2 was best described by a multiple linear regression with terms for adult abundance, TP, and predation. Mean recruitment of age-2 fish in the 1978–1998 year-classes predicted by using the two models in sequence was only about 20% greater than the observed mean recruitment. Model estimates fit the measured data exceptionally well for all but the largest four year-classes, which suggests that the models will facilitate improvement in estimates of trophic transfer due to alewives.

Transactions of the American Fisheries Society

Designing mark-recapture studies to reduce effects of distance weighting on movement distance distributions of stream fishes

Mark-recapture studies generate biased, or distance-weighted, movement data because short distances are sampled more frequently than long distances. Using models and field data, we determined how study design affects distance weighting and the movement distributions of stream fishes. We first modeled distance weighting as a function of recapture section length in an unbranching stream. The addition of an unsampled tributary to one of these models substantially increased distance weighting by decreasing the percentage of upstream distances that were sampled. Similarly, the presence of multiple tributaries in the field study resulted in severe bias. However, increasing recapture section length strongly affected distance weighting in both the model and the field study, producing a zone where the number of fish moving could be estimated with little bias. Subsampled data from the field study indicated that longer median (three of three species) and maximum distances (two of three species) can be detected by increasing the length of the recapture section. The effect was extreme for bluehead chub Nocomis leptocephalus, a highly mobile species, which exhibited a longer median distance (133 m versus 60 m), a longer maximum distance (1,144 m versus 708 m), and a distance distribution that differed in shape when the full (4,123-m recapture section) and subsampled (1,978-m recapture section) data sets were compared. Correction factors that adjust the observed number of movements to undersampled distances upwards and those to oversampled distances downwards could not mitigate the distance weighting imposed by the shorter recapture section. Future studies should identify the spatial scale over which movements can be accurately measured before data are collected. Increasing recapture section length a priori is far superior to using post hoc correction factors to reduce the influence of distance weighting on observed distributions. Implementing these strategies will be especially important in stream networks where fish can follow multiple pathways out of the recapture section.

Transactions of the American Fisheries Society

Effects of lowhead dams on riffle-dwelling fishes and macroinvertebrates in a Midwestern river

Many studies have assessed the effects of large dams on fishes and macroinvertebrates, but few have examined the effects of lowhead dams. We sampled fishes, macroinvertebrates, habitat, and physicochemistry monthly from November 2000 to October 2001 at eight gravel bar sites centered around two lowhead dams on the Neosho River, Kansas. Sites included a reference site and a treatment site both upstream and downstream from each dam. Multivariate analysis of variance indicated that habitat, but not physicochemistry, varied immediately upstream and down-stream from the dams, with resultant effects on macroinvertebrate and fish assemblages. Compared with reference sites, upstream treatment sites were deeper and had lower velocities and downstream treatment sites were shallower and had higher velocities; both upstream and downstream treatment sites had greater substrate compaction than reference sites. Macroinvertebrate richness did not differ among site types, but abundance was lowest at downstream treatment sites and evenness was lowest at upstream treatment sites. Fish species richness did not differ among site types, but abundance was highest at downstream reference sites and evenness was highest at upstream sites. The abundance of some benthic fishes was influenced by the dams, including that of the Neosho madtom Noturus placidus, which was lowest immediately upstream and downstream from dams, and those of the suckermouth minnow Phenacobius mirabilis, orangethroat darter Etheostoma spectabile, and slenderhead darter Percina phoxocephala, which were highest in downstream treatment sites. Although limited to one system during a 1-year period, this study suggests that the effects of lowhead dams on fishes, macroinvertebrates, and habitat are similar to those reported for larger dams, providing important considerations for riverine ecosystem conservation efforts.

Transactions of the American Fisheries Society

Spatial variation in fish species richness of the upper Mississippi River system

Important natural environmental gradients, including the connectivity of off-channel aquatic habitats to the main-stem river, have been lost in many reaches of the upper Mississippi River system, and an understanding of the consequences of this isolation is lacking in regard to native fish communities. The objectives of this study were to describe patterns of fish species richness, evenness, and diversity among representative habitats and river reaches and to examine the relationship between fish species richness and habitat diversity. Each year (1994-1999) fish communities of main-channel borders (MCB), side channel borders (SCB), and contiguous backwater shorelines (BWS) were sampled using boat-mounted electrofishing, mini-fyke-nets, tyke nets, hoop nets, and seines at a standardized number of sites. A total of 0.65 million fish were collected, representing 106 species from upper Mississippi River Pools 4, 8, 13, and 26; the open (unimpounded) river reach; and the La Grange Reach of the Illinois River. Within pools, species richness based on rarefaction differed significantly among habitats and was highest in BWS and lowest in MCB (P < 0.0001). At the reach scale, Pools 4, 8, and 13 consistently had the highest species richness and Pool 26, the open-river reach, and the La Grange Reach were significantly lower (P < 0.0001). Species evenness and diversity indices showed similar trends. The relationship between native fish species richness and habitat diversity was highly significant (r(2) = 0.85; P = 0.0091). These results support efforts aimed at the conservation and enhancement of connected side channels and backwaters. Although constrained by dams, pools with high native species richness could serve as a relative reference. The remnants of natural riverine dynamics that remain in these reaches should be preserved and enhanced; conditions could be used to guide restoration activities in more degraded reaches.

Illinois, Iowa, Minnesota, Missouri, Wisconsin

Do predators influence the distribution of age-0 kokanee in a Colorado Reservoir?

Seasonal changes in reservoir conditions such as productivity, light, and temperature create spatiotemporal variation in habitat that may segregate or aggregate predators and prey, producing implications for the distribution, growth, and survival of fishes. We used hydroacoustics to document the diel vertical distribution of age-0 kokanee Oncorhynchus nerka relative to environmental gradients at Blue Mesa Reservoir, Colorado, during May-August of 2002. Temperature, light, and zooplankton density profiles were examined relative to foraging conditions for kokanee and their primary predator, lake trout Salvelinus namaycush. Age-0 kokanee displayed large diel vertical migrations in May despite the lack of an energetic advantage before reservoir stratification. Age-0 kokanee minimized near-surface foraging at this time, perhaps to avoid predation by visual predators, such as lake trout, in the well-lit surface waters. Strong reservoir stratification in midsummer appeared to provide a thermal refuge from lake trout that the kokanee exploited. By August vertical migrations were shallow and most kokanee remained in the epilimnion throughout the day. Although the energetic implications of the late-summer strategy are unclear, it appears that kokanee were responding to changes in their predator environment. A robust model for kokanee diel vertical migration across a range of systems should include a predator avoidance component.

Colorado

Overwinter survival of juvenile lake herring in relation to body size, physiological condition, energy stores, and food ration

Populations of lake herring Coregonus artedi in Lake Superior have exhibited high recruitment variability over the past three decades. To improve our understanding of the mechanisms which influence year-class strength, we conducted a 225-d laboratory experiment to evaluate the effects of body size, physiological condition, energy stores, and food ration on the winter survival of age-0 lake herring. Small (total length (TL) range = 60&ndash;85 mm) and large (TL range = 86&ndash;110 mm) fish were maintained under thermal and photoperiod regimes that mimicked those in Lake Superior from October through May. Fish in each size-class were maintained at two feeding treatments: brine shrimp Artemia spp. ad libitum and no food. The mortality of large lake herring (fed, 3.8%; starved, 20.1%) was significantly less than that of small fish (fed, 11.7%; starved, 32.0%). Body condition and crude lipid content declined for all fish over the experiment; however, these variables were significantly greater for large fed (0.68% and 9.8%) and small fed (0.65% and 7.3%) fish than large starved (0.49% and 5.7%) and small starved (0.45% and 4.8%) individuals. Final crude protein and gross energy contents were also significantly greater in large fed lake herring (17.6% and 1,966 cal/g), followed by small fed (17.1% and 1,497 cal/g), large starved (15.4% and 1,125 cal/g), and small starved (13.2% and 799 cal/g) fish. Lake herring that died during the experiment had significantly lower body condition and energy stores relative to those of the surviving fish. These results suggest that the depletion of energy stores contributes to greater winter mortality of small lake herring with limited energy uptake and may partially explain the variability in recruitment observed in Lake Superior.

Transactions of the American Fisheries Society

Home range and space use patterns of flathead catfish during the summer-fall period in two Missouri streams

Flathead catfish Pylodictis olivaris were radio-tracked in the Grand River and Cuivre River, Missouri, from late July until they moved to overwintering habitats in late October. Fish moved within a definable area, and although occasional long-distance movements occurred, the fish typically returned to the previously occupied area. Seasonal home range was calculated with the use of kernel density estimation, which can be interpreted as a probabilistic utilization distribution that documents the internal structure of the estimate by delineating portions of the range that was used a specified percentage of the time. A traditional linear range also was reported. Most flathead catfish (89%) had one 50% kernel-estimated core area, whereas 11% of the fish split their time between two core areas. Core areas were typically in the middle of the 90% kernel-estimated home range (58%), although several had core areas in upstream (26%) and downstream (16%) portions of the home range. Home-range size did not differ based on river, sex, or size and was highly variable among individuals. The median 95% kernel estimate was 1,085 m (range, 70&ndash; 69,090 m) for all fish. The median 50% kernel-estimated core area was 135 m (10&ndash;2,260 m). The median linear range was 3,510 m (150&ndash;50,400 m). Fish pairs with core areas in the same and neighboring pools had static joint space use values of up to 49% (area of intersection index), indicating substantial overlap and use of the same area. However, all fish pairs had low dynamic joint space use values (<0.07; coefficient of association), indicating that fish pairs were temporally segregated, rarely occurring in the same location at the same time.

Missouri