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A central goal of population ecology is to establish linkages between life history strategy, disturbance, and population dynamics. Globally, disturbance events such as drought and invasive species have dramatically impacted stream fish populations and contributed to sharp declines in freshwater biodiversity. Here, we used RAMAS Metapop to construct stage-based demographic metapopulation models for stream fishes with periodic, opportunistic, and equilibrium life history strategies and assessed their responses to the effects of invasion (reduced carrying capacity), extended drought (reduced survival and fecundity), and the combined effects of both disturbances. Our models indicated that populations respond differentially to disturbance based on life history strategy. Equilibrium strategists were best able to deal with simulated invasion. Periodic strategists did well under lower levels of drought, whereas opportunistic strategists outperformed other life histories under extreme seasonal drought. When we modeled additive effects scenarios, these disturbances interacted synergistically, dramatically increasing terminal extinction risk for all three life history strategies. Modeling exercises that examine broad life history categories can help to answer fundamental ecological questions about the relationship between disturbance resilience and life history, as well as help managers to develop generalized conservation strategies when species-specific data are lacking. Our results indicate that life history strategy is a fundamental determinant of population trajectories, and that disturbances can interact synergistically to dramatically impact extinction outcomes.

Hydrologic variation can play a major role in structuring stream fish assemblages and relationships between hydrology and biology are likely to be influenced by flow regime. We hypothesized that more variable flow regimes would have lower and more variable species richness, higher species turnover and lower assemblage stability, and greater abiotic environment-fish relationships than more stable flow regimes. We sampled habitats (pool, run, and riffle) in three Runoff/Intermittent Flashy streams (highly variable flow regime) and three Groundwater Flashy streams (less variable flow regime) seasonally (spring, early summer, summer and autumn) in 2002 (drought year) and 2003 (wet year). We used backpack electrofishing and three-pass removal techniques to estimate fish species richness, abundance and density. Fish species richness and abundance remained relatively stable within streams and across seasons, but densities changed substantially as a result of decreased habitat volume. Mixed model analysis showed weak response variable-habitat relationships with strong season effects in 2002, and stronger habitat relationships and no season effect in 2003, and flow regime was not important in structuring these relationships. Seasonal fish species turnover was significantly greater in 2002 than 2003, but did not differ between flow regimes. Fish assemblage stability was significantly lower in Runoff/Intermittent Flashy than Groundwater Flashy streams in 2002, but did not differ between flow regimes in 2003. Redundancy analysis showed fish species densities were well separated by flow regime in both years. Periodic and opportunistic species were characteristic of Runoff/Intermittent Flashy streams, whereas mainly equilibrium species were characteristic of Groundwater Flashy streams. We found that spatial and temporal variation in hydrology had a strong influence on fish assemblage dynamics in Ozark streams with lower assemblage stability and greater fluctuations in density in more hydrologically variable streams and years. Understanding relationships between fish assemblage structure and hydrologic variation is vital for conservation of fish biodiversity. Future work should consider addressing how alteration of hydrologic variation will affect biotic assemblages.

Aquatic organisms have adapted over evolutionary time-scales to hydrologic variability represented by the natural flow regime of rivers and streams in their unimpaired state. Rapid landscape change coupled with growing human demand for water have altered natural flow regimes of many rivers and streams on a global scale. Climate non-stationarity is expected to further intensify hydrologic variability, placing increased pressure on aquatic communities. Using a machine learning approach and georeferenced species occurrence data, we modeled and mapped spatial patterns of hydrologic disturbance for streams in Arkansas, Missouri, and eastern Oklahoma. Random forest (RF) models trained on fish community data, hydrologic, and landscape metrics for gaged streams in the National Hydrography (NHDPlusV2) database were used to predict a hydrologic disturbance index (HDI) for ungaged streams. The HDI is part of the USGS Geospatial Attributes of Gages for Evaluating Streamflow (GAGESII) database and is a composite index of watershed-scale disturbance from anthropogenic stressors. Fish presence/absence data had similar overall model prediction accuracy (77%; 95% CI: 0.74, 0.80) as flow variables (76%; CI: 0.73, 0.80). Including topographic variables increased the RF prediction accuracy of both the fish (90%; CI: 0.88, 0.92) and flow models (86%; CI: 0.84, 0.89). Spatial patterns of hydrologic disturbance suggest distinct ecohydrological regions exist where conservation actions may be focused. Streams with low HDI were predominately located in the Ozark Highlands, Boston Mountains, and Ouachita Mountains. Correlation analysis of HDI by flow regime showed groundwater stable streams had the lowest disturbance frequency, with over 50% of stream reaches with low HDI located in forested land cover. HDI was highest for big rivers, intermittent runoff streams and streams in areas of agricultural land use. Our results show long-term georeferenced biological data can provide a valuable resource for predictive modeling of hydrologic disturbance for ungaged rivers and streams.

Ozark Bass (Ambloplites constellatus) is an understudied, endemic fish species in the Upper White River Basin of northern Arkansas. This study was part of an effort by fisheries managers to gather baseline data about the Ozark Bass to aid in understanding population dynamics and contribute to the limited data available for use in determining the efficacy of harvest regulations. Select population characteristics of Ozark Bass in two northern Arkansas streams were determined, population characteristics of Ozark Bass were compared to Shadow Bass (Ambloplites ariommus) and Rock Bass (Ambloplites rupestris) data collected from previous studies in southern Missouri, and relative condition, length-at-age, and annual survival of Ozark Bass were compared between sample streams. Sampling occurred in Crooked Creek and the Buffalo River during summer 2013 via boat electroshocking. Length and weight data were recorded for all Ozark Bass collected, and fish ages were determined through selective otolith retrieval and age-length keys. Ozark Bass in Crooked Creek had greater relative condition than Ozark Bass in Buffalo River (P 0.05) between sexes for fish collected from only the Buffalo River. Ozark Bass mean annual survival was similar between Crooked Creek (55% ± 5% as 95% confidence interval (CI)) and the Buffalo River (50% ± 7% CI) for fish age 2 to 9. Calculated Ozark Bass lengths-at-age for fish from both streams were comparable to the Von Bertalanffy growth estimates, except the Buffalo River age 7 category where there was only one observation. The relationship between Ozark Bass age and length differed between sampled streams, and variability in growth rates and length-at-age were observed among Ambloplites species. Results of this study contribute to the understanding of the population dynamics of the Ozark Bass that will lead to improved fisheries management.

Drought and nutrient pollution can affect the dynamics of stream ecosystems in diverse ways. While the individual effects of both stressors are broadly examined in the literature, we still know relatively little about if and how these stressors interact. Here, we performed a mesocosm experiment that explores the compounded effects of seasonal drought via water withdrawals and nutrient pollution (1.0 mg/L of N and 0.1 mg/L of P) on a subset of Ozark stream community fauna and ecosystem processes. We observed biological responses to individual stressors as well as both synergistic and antagonistic stressor interactions. We found that drying negatively affected periphyton assemblages, macroinvertebrate colonization, and leaf litter decomposition in shallow habitats. However, in deep habitats, drought-based increases in fish density caused trophic cascades that released algal communities from grazing pressures; while nutrient enrichment caused bottom-up cascades that influenced periphyton variables and crayfish growth rates. Finally, the combined effects of drought and nutrient enrichment interacted antagonistically to increase survival in longear sunfish; and stressors acted synergistically on grazers causing a trophic cascade that increased periphyton variables. Because stressors can directly and indirectly impact biota—and that the same stressor pairing can act differentially on various portions of the community simultaneously—our broad understanding of individual stressors might not adequately inform our knowledge of multi-stressor systems.

The natural flow regime is considered the “master variable” in lotic systems, controlling structure and function at organismal, population, community, and ecosystem levels. We sought to estimate forested headwater stream metabolism across two dominant flow regimes (Runoff and Groundwater) in northern Arkansas and evaluate potential differences in, and drivers of, gross primary production, ecosystem respiration, and net ecosystem metabolism. Flow regimes differed in intermittency, substrate heterogeneity, hyporheic connectivity, and dominant water source (subsurface runoff vs. groundwater), which we expected to result in differences in primary production and respiration. Average daily gross primary production (GPP) and ecosystem respiration (ER) estimated from field data collected from May 2015-June 2016 tended to be greater in Groundwater streams. Respiration was positively related to discharge (R2= 0.98 p< 0.0001) and net metabolism became more heterotrophic with increasing average annual discharge across sites (R2= 0.94, p= 0.0008). Characterizing ecosystem-level responses to differences in flow can reveal mechanisms governing stream metabolism and, in turn, provide information regarding trophic state and energy inputs as efforts continue to determine global trends in aquatic carbon sources and fates.

Given the importance of crayfish in stream ecosystems, gaining insight into the role of stream permanence in maintaining predator–prey interactions is critical. Our objectives were to determine the influence of stream permanence and season on crayfish predation and assess the role of stream permanence and crayfish density on the presence of predators, while accounting for imperfect detection. We conducted surveys of crayfish density, mammalian scat, and environmental variables within 10 intermittent and 10 permanent streams in the Ozark Highlands. We used occupancy modeling to assess the relationship between predator presence, crayfish density, and environmental variables. Stream permanence did not play a role in determining relative frequency of occurrence or percent volume of crayfish prey in mammalian diets. However, percent volume and relative frequency of crayfish prey found in scats differed by season, with both highest in spring and summer. The relative frequency and percent volume of fish prey showed a significant interaction of season by stream permanence, which may be the first instance of this observation. Procyon lotor (Raccoon) had the highest detection probability (p = 0.39), whereas Neovison vison (American Mink; p = 0.15) and Lontra canadensis (River Otter; p = 0.03) had low detection probabilities. Further study into predator–prey interactions in the context of hydrology, particularly when related to imperiled groups like freshwater crayfishes, is needed since climate change is expected to alter hydrologic patterns.

Crayfish are ecologically important in freshwater systems worldwide and are imperiled in North America and globally. We sought to examine landscape- to local-scale environmental variables related to occupancy and detection probability of a suite of stream-dwelling crayfish species. We used a quantitative kickseine method to sample crayfish presence at 102 perennial stream sites with eight surveys per site. We modeled occupancy (psi) and detection probability (P) and local- and landscape-scale environmental covariates. We developed a set of a priori candidate models for each species and ranked models using (Q)AICc. Detection probabilities and occupancy estimates differed among crayfish species with Orconectes eupunctus, O. marchandi, and Cambarus hubbsi being relatively rare (psi 0.60) with high detection probability (0.81). Detection probability was often related to local habitat variables current velocity, depth, or substrate size. Important environmental variables for crayfish occupancy were species dependent but were mainly landscape variables such as stream order, geology, slope, topography, and land use. Landscape variables strongly influenced crayfish occupancy and should be considered in future studies and conservation plans.

Fishes often exhibit phenotypic divergence across gradients of abiotic and biotic selective pressures. In streams, many of the known selective pressures driving phenotypic differentiation are largely influenced by hydrologic regimes. Because flow regimes drive so many attributes of lotic systems, we hypothesized fish exhibit phenotypic divergence among streams with different flow regimes. We used a comparative field study to investigate the morphological divergence of Campostoma anomalom (central stonerollers) among streams characterized by highly variable, intermittent flow regimes and streams characterized by relatively stable, groundwater flow regimes. We also conducted a mesocosm experiment to compare the plastic effects of one component of flow regimes, water velocity, on morphology of fish from different flow regimes. We observed differences in shape between flow regimes likely driven by differences in allometric growth patterns. Although we observed differences in morphology across flow regimes in the field, C. anomalum did not exhibit morphologic plasticity in response to water velocity alone. This study contributes to the understanding of how complex environmental factors drive phenotypic divergence and may provide insight into the evolutionary consequences of disrupting natural hydrologic patterns, which are increasingly threatened by climate change and anthropogenic alterations.

Restrictive angling regulations in tailwater trout fisheries may be unsuccessful if food availability limits energy for fish to grow. We examined spatial and temporal variation in energy intake and growth in populations of Brown Trout Salmo trutta and Rainbow Trout Oncorhynchus mykiss within three catch-and-release (C-R) areas in Arkansas tailwaters to evaluate food availability compared with consumption. Based on bioenergetic simulations, Rainbow Trout fed at submaintenance levels in both size-classes (≤400 mm TL, >400 mm TL) throughout most seasons. A particular bottleneck in food availability occurred in the winter for Rainbow Trout when the daily ration was substantially below the minimum required for maintenance, despite reduced metabolic costs associated with lower water temperatures. Rainbow Trout growth rates followed a similar pattern to consumption with negative growth rates during the winter periods. All three size-classes ( 400 mm TL) of Brown Trout experienc...

A central goal of population ecology is to establish linkages between life history strategy, disturbance, and population dynamics. Globally, disturbance events such as drought and invasive species have dramatically impacted stream fish populations and contributed to sharp declines in freshwater biodiversity. Here, we used RAMAS Metapop to construct stage-based demographic metapopulation models for stream fishes with periodic, opportunistic, and equilibrium life history strategies and assessed their responses to the effects of invasion (reduced carrying capacity), extended drought (reduced survival and fecundity), and the combined effects of both disturbances. Our models indicated that populations respond differentially to disturbance based on life history strategy. Equilibrium strategists were best able to deal with simulated invasion. Periodic strategists did well under lower levels of drought, whereas opportunistic strategists outperformed other life histories under extreme seasonal drought. When we modeled additive effects scenarios, these disturbances interacted synergistically, dramatically increasing terminal extinction risk for all three life history strategies. Modeling exercises that examine broad life history categories can help to answer fundamental ecological questions about the relationship between disturbance resilience and life history, as well as help managers to develop generalized conservation strategies when species-specific data are lacking. Our results indicate that life history strategy is a fundamental determinant of population trajectories, and that disturbances can interact synergistically to dramatically impact extinction outcomes.

Temperature increases due to climate change over the coming century will likely affect smallmouth bass (Micropterus dolomieu) growth in lotic systems at the southern extent of their native range. However, the thermal response of a stream to warming climate conditions could be affected by the flow regime of each stream, mitigating the effects on smallmouth bass populations. We developed bioenergetics models to compare change in smallmouth bass growth rate potential (GRP) from present to future projected monthly stream temperatures across two flow regimes: runoff and groundwater-dominated. Seasonal differences in GRP between stream types were then compared. The models were developed for fourteen streams within the Ozark–Ouachita Interior Highlands in Arkansas, Oklahoma and Missouri, USA, which contain smallmouth bass. In our simulations, smallmouth bass mean GRP during summer months decreased by 0.005 g g−1 day−1 in runoff streams and 0.002 g g−1 day−1 in groundwater streams by the end of century. Mean GRP during winter, fall and early spring increased under future climate conditions within both stream types (e.g., 0.00019 g g−1 day−1 in runoff and 0.0014 g g−1 day−1 in groundwater streams in spring months). We found significant differences in change in GRP between runoff and groundwater streams in three seasons in end-of-century simulations (spring, summer and fall). Potential differences in stream temperature across flow regimes could be an important habitat component to consider when investigating effects of climate change as fishes from various flow regimes that are relatively close geographically could be affected differently by warming climate conditions.

Natural disturbance is an integral component of most ecosystems and occurs in 3 different forms: pulse, press, and ramp. In lotic ecosystems, seasonal drought is a major form of disturbance, particularly in intermittent headwater streams, which often are reduced to pools that serve as refuges for biota. We used simulated intermittent stream pools to compare the effects of control, pulse, and press drying on growth and survival in 3 fish species (Lepomis megalotis, Campostoma anomalum, and Etheostoma spectabile) commonly found together in drought-prone streams in the Ozark Highlands, USA. We also compared effects on benthic community structure, including periphyton and chironomid density and sediment in deep (permanently watered) and shallow (intermittently dewatered) habitat. Only one species, L. megalotis, showed a significant reduction in length and mass growth in press drying compared with control treatments. Drying and type of drying had no effect on survival of any fish species. Drying and ty...

We examined flow alteration-ecology relationships in benthic macroinvertebrate, fish, and crayfish assemblages in Ozark Highland streams, USA, over two years with contrasting environmental conditions, a drought year (2012) and a flood year (2013). We hypothesized that: 1) there would be temporal variation in flow alteration-ecology relationships between the two years, 2) flow alteration-ecology relationships would be stronger during the drought year vs the flood year, and 3) fish assemblages would show the strongest relationships with flow alteration. We used a quantitative richest-targeted habitat (RTH) method and a qualitative multi-habitat (QMH) method to collect macroinvertebrates at 16 USGS gaged sites during both years. We used backpack electrofishing to sample fish and crayfish at 17 sites in 2012 and 11 sites in 2013. We used redundancy analysis to relate biological response metrics, including richness, diversity, density, and community-based metrics, to flow alteration. We found temporal variation in flow alteration-ecology relationships for all taxa, and that relationships differed greatly between assemblages. We found relationships were stronger for macroinvertebrates during the drought year but not for other assemblages, and that fish assemblage relationships were not stronger than the invertebrate taxa. Magnitude of average flow, frequency of high flow, magnitude of high flow, and duration of high flow were the most important categories of flow alteration metrics across taxa. Alteration of high and average flows was more important than alteration of low flows. Of 32 important flow alteration metrics across years and assemblages, 19 were significantly altered relative to expected values. Ecological responses differed substantially between drought and flood years, and this is likely to be exacerbated with predicted climate change scenarios. Differences in flow alteration-ecology relationships among taxonomic groups and temporal variation in relationships illustrate that a complex suite of variables should be considered for effective conservation of stream communities related to flow alteration.

Crayfish are abundant and important consumers in aquatic food webs and crayfish invasions have demonstrated strong effects of crayfish on multiple trophic levels. Density may be an important factor determining the role of omnivorous crayfish in benthic communities, especially if density alters the strength of trophic interactions. The effect of crayfish density on a simple benthic food web using ceramic tiles was examined in three treatments (crayfish exclusion cage, cage control (open to crayfish), and exposed ceramic tiles) in mesocosms stocked with 6, 12, or 18 crayfish·m-2. We hypothesized that at low densities crayfish consumption of herbivorous chironomids would increase algal abundance, but at high densities crayfish would reduce both periphyton and invertebrates. In the experiment, periphyton and chironomid abundance increased with declining crayfish biomass on day 30 but not day 15. The magnitude of crayfish effects on day 15 periphyton chlorophyll a abundance increased with crayfish biomass, but crayfish effects on day 30 periphyton chlorophyll a or chironomid biomass did not increase with crayfish biomass. In this experiment there was little evidence for a trophic cascade at low crayfish densities and strong omnivory by crayfish dominated trophic dynamics.

We examined trophic dynamics of a stream food web where invasive Orconectes neglectus appear to be displacing native O. eupunctus in the Spring River drainage of the Ozark Highlands, Missouri and Arkansas, USA. We collected crayfish species and possible food sources seasonally from a site of sympatry on the South Fork Spring River. We determined diet overlap and potential for competition between O. eupunctus and O. neglectus, and investigated seasonal variation using carbon and nitrogen stable isotope analyses and gut content analyses. Gut content analysis showed both species of crayfish consumed mainly detritus during summer and spring, with other prey categories varying by species and season. Stable isotope analysis showed that O. eupunctus and O. neglectus relied on invertebrates as a major energy and nutrient source throughout summer, autumn, and spring, and the two species showed differences in their stable isotope signatures during spring and summer, but not autumn. Given the trophic overlap between O. eupunctus and O. neglectus, there is a potential for the two species to compete for food and to be ecologically redundant. Ecological redundancy can lead to reduced effects on ecosystem function post-invasion, and therefore examining ecological redundancy of potential invaders should be a conservation priority.

The Yellowcheek Darter (Etheostoma moorei) is a rare fish endemic to the Little Red River watershed in the Boston Mountains of northern Arkansas. Remaining populations of this species are geographically isolated and declining, and the species was listed in 2011 as federally endangered. Populations have declined, in part, due to intense seasonal stream drying and inundation of lower reaches by a reservoir. We used a kick seine sampling approach to examine distribution and abundance of Yellowcheek Darter populations in the Middle Fork and South Fork Little Red River. We used presence data to estimate occupancy rates and detection probability and examined relationships between Yellowcheek Darter density and environmental variables. The species was found at five Middle Fork and South Fork sites where it had previously been present in 2003–2004. Occupancy rates were >0.6 but with wide 95% CI, and where the darters occurred, densities were typical of other Ozark darters but highly variable. Detection probabilit...

Natural flow regimes represent the hydrologic conditions to which native aquatic organisms are best adapted. We completed a regional river classification and quantitative descriptions of each natural flow regime for the Ozark–Ouachita Interior Highlands region of Arkansas, Missouri and Oklahoma. On the basis of daily flow records from 64 reference streams, seven natural flow regimes were identified with mixture model cluster analysis: Groundwater Stable, Groundwater, Groundwater Flashy, Perennial Runoff, Runoff Flashy, Intermittent Runoff and Intermittent Flashy. Sets of flow metrics were selected that best quantified nine ecologically important components of these natural flow regimes. An uncertainty analysis was performed to avoid selecting metrics strongly affected by measurement uncertainty that can result from short periods of record. Measurement uncertainties (bias, precision and accuracy) were assessed for 170 commonly used flow metrics. The ranges of variability expected for select flow metrics under natural conditions were quantified for each flow regime to provide a reference for future assessments of hydrologic alteration. A random forest model was used to predict the natural flow regimes of all stream segments in the study area based on climate and catchment characteristics, and a map was produced. The geographic distribution of flow regimes suggested distinct ecohydrological regions that may be useful for conservation planning. This project provides a hydrologic foundation for future examination of flow–ecology relationships in the region. Published 2014. This article is a U.S. Government work and is in the public domain in the USA.

SUMMARY 1. Crayfishes and other freshwater aquatic fauna are particularly at risk globally due to anthropogenic demand, manipulation and exploitation of freshwater resources and yet are often understudied. The Ozark faunal region of Missouri and Arkansas harbours a high level of aquatic biological diversity, especially in regard to endemic crayfishes. Three such endemics, Orconectes eupunctus, Orconectes marchandi and Cambarus hubbsi, are threatened by limited natural distribution and the invasions of Orconectes neglectus. 2. We examined how natural and anthropogenic abiotic factors influence these three species across multiple spatial scales. Local and landscape environmental variables were used as predictors in classification and regression tree models at stream segment and segmentshed scales to determine their relation to presence/absence and density of the three species. 3. Orconectes eupunctus presence was positively associated with stream size, current velocity and spring flow volume. Orconectes marchandi presence was predicted primarily by dolomite geology and water chemistry variables. Cambarus hubbsi was associated with larger stream size, with highest densities occurring in deep waters. Stream segment and segmentshed scale models were similar, but there were important differences based on species and response variables (presence/absence versus density). Stream segment scale models consistently performed better than or equal to segmentshed scale models. 4. Anthropogenic abiotic environmental variables were of minor importance in most models, with the exception of O. marchandi being negatively related to road density and human population density. Classification tree models predicting distribution performed well when compared to random assignment, but regression trees were generally poor in explaining variation in density. 5. We found that a range of environmental variables were important in predicting crayfish distribution and abundance at multiple spatial scales and their importance was species-, response variable- and scale dependent. We would encourage others to examine the influence of spatial scale on species distribution and abundance patterns.

Predation from aquatic and terrestrial predators are important factors structuring the size and depth distribution of aquatic prey. We conducted mesocosm and tethering experiments on Little Mulberry Creek in northwest Arkansas during low flows to examine the effects of predators on fish and crayfish survival in intermittent streams. Using shallow artificial pools (10 cm deep) and predator exclusions, we tested the hypothesis that large-bodied fish are at greater risk from terrestrial predators in shallow habitats compared to small-bodied individuals. Twenty-four circular pools (12 open top, 12 closed top) were stocked with two size classes of Campostoma anomalum (Central Stoneroller) and deployed systematically in a single stream pool. In addition, we used a crayfish tethering experiment to test the hypothesis that the survival of small and large crayfish is greater in shallow and deep habitats, respectively. We tethered two size classes of Orconectes meeki meeki (Meek's Crayfish) along shallow and deep transects in two adjacent stream pools and measured survival for 15 days. During both experiments, we monitored the presence or absence of predators by visual observation and from scat surveys. We demonstrated a negative effect of terrestrial predators on Central Stoneroller survival in the artificial pools, and larger individuals were more susceptible to predation. In contrast, small crayfish experienced low survival at all depths and large crayfish were preyed upon much less intensively during the tethering study, particularly in the pool with larger substrate. More studies are needed to understand how stream drying and environmental heterogeneity influence the complex interactions between predator and prey populations in intermittent streams.

The Big Creek Crayfish, Orconectes peruncus, is native to the St. Francis River drainage in Missouri, USA and is often absent where the introduced Woodland Crayfish, Orconectes hylas, has established. We performed a field experiment to determine whether effects of current abiotic conditions and interspecific competition with O. hylas were responsible for displacement of O. peruncus from parts of their former range. We examined growth and survival of juvenile male O. peruncus exposed to juvenile male O. hylas in enclosures at two sites in the former range of O. peruncus. Enclosures contained 8 (low density) or 16 individuals (high density) and had O. peruncus only (control) or both species (interspecific treatment). Juvenile O. peruncus were able to survive and grow in portions of their former range, implicating biotic versus abiotic factors in the displacement of O. peruncus. Survival rates of O. peruncus did not differ among treatments at either site. Orconectes peruncus showed significant growth in all treatments and interspecific effects were not greater than intraspecific effects on O. peruncus growth rates. High-density treatments showed significantly reduced O. peruncus growth rates compared to low-density treatments, except in Carver Creek interspecific treatments. When considered in the context of previous studies examining the effects of O. hylas on O. peruncus, results suggest that neither direct competition between juvenile males of the two species or abiotic change are responsible for the decreased range of O. peruncus. Additional research is required to determine the mechanism(s) driving the displacement of O. peruncus.

Drought and summer drying can be important disturbance events in many small streams leading to intermittent or isolated habitats. We examined what habitats act as refuges for fishes during summer drying, hypothesizing that pools would act as refuge habitats. We predicted that during drying fish would show directional movement into pools from riffle habitats, survival rates would be greater in pools than in riffles, and fish abundance would increase in pool habitats. We examined movement, survival and abundance of three minnow species, bigeye shiner (Notropis boops), highland stoneroller (Campostoma spadiceum) and creek chub (Semotilus atromaculatus), during seasonal stream drying in an Ozark stream using a closed robust multi-strata mark-recapture sampling. Population parameters were estimated using plausible models within program MARK, where a priori models are ranked using Akaike’s Information Criterion. Creek chub showed directional movement into pools and increased survival and abundance in pools during drying. Highland stonerollers showed strong directional movement into pools and abundance increased in pools during drying, but survival rates were not significantly greater in pools than riffles. Bigeye shiners showed high movement rates during drying, but the movement was non-directional, and survival rates were greater in riffles than pools. Therefore, creek chub supported our hypothesis and pools appear to act as refuge habitats for this species, whereas highland stonerollers partly supported the hypothesis and bigeye shiners did not support the pool refuge hypothesis. Refuge habitats during drying are species dependent. An urgent need exists to further understand refuge habitats in streams given projected changes in climate and continued alteration of hydrological regimes.

River otters ( Lontra canadensis ) are important predators in aquatic ecosystems, but few studies quantify their prey consumption. We trapped crayfish monthly as an index of availability and collected otter scat for diet analysis in the Ozark Mountains of northwestern Arkansas, USA. We measured otter daily energy expenditure (DEE) with the doubly labeled water method to develop a bioenergetics model for estimating monthly prey consumption. Meek’s crayfish ( Orconectes meeki ) catch-per-unit-effort was positively related to stream temperature, indicating that crayfish were more available during warmer months. The percentage frequency of occurrence for crayfish in scat samples peaked at 85.0% in summer and was lowest (42.3%) in winter. In contrast, the percentage occurrence of fish was 13.3% in summer and 57.7% in winter. Estimates of DEE averaged 4738 kJ·day–1 for an otter with a body mass of 7842 g. Total biomass consumption ranged from 35 079 to 52 653 g·month–1 (wet mass), corresponding to a high proportion of fish and crayfish in the diet, respectively. Otter consumption represents a large fraction of prey production, indicating potentially strong effects of otters on trophic dynamics in stream ecosystems.

Climate change will affect stream systems in numerous ways over the coming century. Globally, streams are expected to experience changes in temperature and flow regime. Previous work has indicated that these changes will likely affect fish distributions, but little work has been conducted examining population level effects of climate change on warmwater fish at the warmest portion of their range. We model several potential climate change-related stressors and the resulting effects on smallmouth bass Micropterus dolomieu populations in the Buffalo National River, Arkansas, USA, located near the southern extent of smallmouth bass range. Smallmouth bass are a popular recreational fish in the region and angler harvest likely contributes substantially to annual mortality. We created a simulation model parameterized with data collected from the Buffalo National River to evaluate the relative importance of climate stressors and angler harvest on smallmouth bass populations. Our simulations suggest that increases in springtime temperature and reductions in river discharge during the spawning period could increase recruitment, resulting in increases in adult abundance (8% higher). However, when increased flooding and drought probabilities are considered, our model indicates the Buffalo National River could experience large reductions in adult smallmouth bass abundance (≥50% decline) and increased probability of extinction compared to present levels. Simulations showed that harvest reduction could be a viable strategy to reduce the negative effects of climate change, but that even with complete closure of harvest, smallmouth bass population levels would still be well below present abundance (46% lower than present). Efforts to reduce flooding and drought effects related to climate change in the Buffalo National River could help offset the predicted reduction in the smallmouth bass population.

Consumers can alter the structure and function of aquatic ecosystems, but the interactions that regulate ecosystem functioning are context dependent. In many streams, environmental disturbance and consumer identity may interact to influence benthic ecosystem structure and function. We predicted that effects of two common consumers (central stonerollers (Campostoma anomalum), and Meek's crayfish (Orconectes meeki meeki)) in Ozark Mountain streams (Arkansas, USA) would be altered by disturbance. In stream mesocosms we crossed disturbance (simulated spate) and consumer identity (crayfish, central stonerollers, or no consumers). Stream ecosystem function was quantified as leafpack decomposition and net primary productivity (NPP). Ecosystem structure was measured as algal chlorophyll-a, ash-free dry mass (AFDM), and chironomid density. Disturbance and interactions between disturbance and consumer identity did not significantly affect any response variable. Chlorophyll-a, AFDM, and chironomid density were influenced by consumer identity, and effects differed between stonerollers and crayfish and with time. Crayfish and stonerollers reduced day 15 chironomid density compared with no consumer treatments. Stonerollers reduced day 30 chironomid density and day 15 periphyton abundance compared with crayfish and no consumer treatments. Crayfish increased leafpack breakdown rates above stoneroller and no consumer treatments, but NPP did not differ among consumer treatments. In this experiment, consumers had stronger effects on benthic structure and function than the disturbance treatment. Future studies should investigate a wider range of disturbance frequency and intensity.

Benthic consumers influence stream ecosystem structure and function, but these interactions depend on environmental context. We experimentally quantified the effects of central stoneroller minnows (Campostoma anomalum (Rafinesque) and Meek’s crayfish (Orconectes meeki meeki (Faxon)) on benthic communities using electric exclusion quadrats in Little Mulberry Creek before (June) and during (August) seasonal stream drying. Unglazed ceramic tiles were deployed in June and August to measure periphyton and invertebrate abundance, and leafpack decomposition and primary production were also measured in August. Relationships between stoneroller and crayfish density and the size of consumer effects were evaluated with multiple linear regression models. Average chlorophyll a abundance was greater on exposed than exclusion tiles in August, but not in June. Sediment dry mass, periphyton ash-free dry mass (AFDM), and chironomid densities on tiles did not differ among treatments in either period. Leaf packs decayed faster in exposed than exclusion treatments (k exposed = 0.038 ± 0.013, k exclusion = 0.007 ± 0.002), but consumer effects were stronger in some pools than others. Leafpack invertebrate biomass and abundance and tile primary productivity did not differ among treatments. Consumer effects on chlorophyll a were related to crayfish and stoneroller density, and effects on chironomid density were related to stoneroller density. These results contrast with a previous exclusion experiment in Little Mulberry Creek that demonstrated strong consumer effects. The influence of stream drying on consumer effects appears to have been reduced by strong spates, underscoring the importance of conducting multi-year studies to determine the magnitude of variability in ecological interactions.

Summary 1. Crayfish are among the most threatened taxa in the world and invasive crayfish are the primary cause of the decline of native crayfish. Most research has emphasised biotic interactions as the mechanism by which native crayfish are displaced by invasives, although crayfish occupy variable environments and the role of disturbance in facilitating crayfish invasion and displacement is understudied. 2. We compared tolerance to a disturbance, stream drying, in a native and invasive crayfish as a potential mechanism to explain their distribution. Our experiments and observations were conducted across scales, from laboratory environmental chambers to stream mesocosms to field sampling. We hypothesised that the invasive crayfish would be more tolerant of desiccation than the native, and that this physiological distinction between the two would be reflected in their distribution in relation to stream drying. 3. In the laboratory, the native crayfish Orconectes eupunctus was less tolerant of desiccation than the invasive Orconectes neglectus chaenodactylus, with all native crayfish dying within 2 days without water, while some of the invasive crayfish survived for nearly 2 weeks. Under simulated stream drying in mesocosms, only the native O. eupunctus survived less well than in a control. Field sampling demonstrated a significant negative relationship between O. eupunctus density and low summer flows, while O. neglectus density was positively associated with low summer flows. The greater resistance of O. neglectus to drying could, through priority effects, inhibit recolonisation by O. eupunctus once flow resumes. 4. Abiotic disturbances are potentially important to the displacement of native by invasive crayfish. Disturbance mediated displacement of aquatic species provides both an opportunity to conserve native species by maintaining or restoring habitat and disturbance regimes and is also a challenge due to increasing human water demand, flow regime alteration and global climate change.

Understanding spatial and temporal variation in the trophic base of stream food webs is critical for predicting population and community stability, and ecosystem function. We used stable isotope ratios (13C/12C, and 15N/14N) to characterize the trophic base of two streams in the Ozark Mountains of northwest Arkansas, U.S.A. We predicted that autochthonous resources would be more important during the spring and summer and allochthonous resources would be more important in the winter due to increased detritus inputs from the riparian zone during autumn leaf drop. We predicted that stream communities would demonstrate increased reliance on autochthonous resources at sites with larger watersheds and greater canopy openness. The study was conducted at three low-order sites in the Mulberry River Drainage (watershed area range: 81–232 km2) seasonally in 2006 and 2007. We used circular statistics to examine community-wide shifts in isotope space among fish and invertebrate consumers in relation to basal resources, including detritus and periphyton. Mixing models were used to quantify the relative contribution of autochthonous and allochthonous energy sources to individual invertebrate consumers. Significant isotopic shifts occurred but results varied by season and site indicating substantial variation in the trophic base of stream food webs. In terms of temporal variation, consumers shifted toward periphyton in the summer during periods of low discharge, but results varied during the interval between summer and winter. Our results did not demonstrate increased reliance on periphyton with increasing watershed area or canopy openness, and detritus was important at all the sites. In our study, riffle–pool geomorphology likely disrupted the expected spatial pattern and stream drying likely impacted the availability and distribution of basal resources.

Despite the prevalence and importance of intermittent streams, few studies have examined their use by crayfishes. We documented persistence in an intermittent stream and use of the hyporheic zone by 2 imperiled crayfishes, Orconectes williamsi and Orconectes meeki meeki, upon stream drying. During 2005 to 2007, we monitored summer flows and crayfish densities in 10 riffles in a Missouri (US) Ozark stream to document stream drying and crayfish persistence. Stream drying variables were riffle wetted area (m2) and current velocity (m/s). Crayfish density estimates were obtained with a quadrat sampler. We sampled monthly (May–August 2006) to examine crayfish response to decreased flows, and excavated dry quadrat samples (to 30 cm depth) after surface flows ceased. Over 2 periods during summer 2007, we used baited traps in the stream and a downstream reservoir where the stream terminates to examine whether crayfish migrated from the stream to permanent water in response to drying. Riffles dried signifi...

The crayfish Orconectes eupunctus is endemic to the Spring and Eleven Point rivers of Arkansas and Missouri, and appears to have been displaced from a portion of its range by the recently introduced crayfish O. neglectus. Life history can be a factor in crayfish species displacements as earlier reproduction or rapid juvenile growth may provide size advantages in competitive interactions, while higher fecundity might permit an invading species to out reproduce a native. We conducted a comparative study to determine if life history differences exist between O. neglectus and O. eupunctus that could contribute to the displacement of O. eupunctus. Crayfish were collected at three sites (each species alone and both species together) monthly from Jul. 2005 through Dec. 2006 by kick-seining. Timing of reproductive events was recorded, fecundity was evaluated through egg counts, and juvenile growth rates were examined using length-frequency histograms. Sex ratios, length-weight relationships, and adult si...

Quantitative sampling of crayfish can be challenging as a result of clustered distributions produced by a variety of factors. We tested the efficiency of a 1-m2 quadrat sampler for estimating the density of riffle-dwelling crayfish. Sampling efficiency was evaluated in an Ozark Plateau stream through a mark–recapture study. We stocked three marked crayfish per square meter into 12 enclosed riffle sections (range, 46–53 m2) and then randomly sampled five 1-m2 quadrats in each of the 12 sections. Sampling efficiency was determined by comparing the estimates of crayfish density to the known density of marked crayfish. We found crayfish sampling efficiency (mean, 69%) to be comparable to that in studies evaluating the quadrat sampler for stream fish, although efficiency across our study riffles ranged widely, from large underestimates (27%) to positively biased overestimates (140%). Coefficients of variation revealed that replicating habitat units within a stream or stream reach may provide more prec...

The coldwater crayfish Orconectes eupunctus is endemic to the Spring and Eleven Point Rivers of Arkansas and Missouri, and appears to have been displaced from a portion of its range by the recently introduced ringed crayfish Orconectes neglectus. We examined competition among juveniles as a potential mechanism for this crayfish species displacement through laboratory and field experiments. Orconectes eupunctus juveniles survived and grew in stream cages in their former range, implicating biotic interactions rather than habitat degradation in the displacement. Laboratory experiments revealed O. neglectus juveniles were dominant in the presence of limited food, whereas size rather than species determined occupancy of limited shelter. In a field competition experiment using stream cages, O. neglectus juveniles did not inhibit growth or reduce survival of O. eupunctus juveniles. Consequently, laboratory evidence of O. neglectus dominance did not correspond with competition under field conditions. Combined with previous studies examining the effects of O. neglectus on O. eupunctus, these results suggest that competition may not be a factor in this crayfish species displacement. Alternate mechanisms for the apparent displacement of O. eupunctus by O. neglectus, such as differential predation or reproductive interference, should be investigated.

Swimming ability of fish species may help explain their persistence or absence in flood-disturbed streams. We used two artificial stream channels with low-complexity (smooth Plexiglas) and high-complexity (rocks glued onto Plexiglas) substrates to determine critical swim speeds (U crit, cm/s) and examine flood resistance behavior of central stonerollers Campostoma anomalum, cardinal shiners Luxilus cardinalis, orangethroat darters Etheostoma spectabile, green sunfish Lepomis cyanellus, and longear sunfish L. megalotis. In the low-complexity treatment, central stonerollers and cardinal shiners had significantly higher U crit than the other species, and orangethroat darters had significantly higher U crit than the sunfish species did. In the high-complexity treatment, the U crit of central stonerollers was significantly higher than that of green sunfish, whereas those for all other species did not differ significantly. Flood resistance behavior was measured using relative velocity (i.e., a fish's c...

We conducted field surveys and experiments to determine whether observed distributions of crayfish among habitats were influenced by differential resource availability, foraging profitability, and predation rates and whether these factors differed with crayfish size and habitat depth. We sampled available food resources (detritus and invertebrates) and shelter as rock substrate in deep (>50 cm) and shallow (

– We evaluated fish assemblage structure during stream drying in north west Arkansas in 2002 and 2003. We sampled fish with a seine and backpack electrofisher and measured habitat variables along transects. Linear regression models were selected a priori and ranked according to Akaike's Information Criterion. In 2002, total fish density was negatively related to pool area and maximum depth, and positively related to canopy openness and substrate diversity. Similarly, central stoneroller (Campostoma anomalum, Rafinesque) and creek chub (Semotilus atromaculatus, Mitchill) densities were negatively related to maximum pool depth. In 2003, fish species richness was positively related to pool area whereas total fish, central stoneroller, and creek chub densities were negatively related to maximum pool depth. In addition, total fish density was negatively related to substrate diversity. Results indicated that physical factors are important predictors of fish assemblage structure during stream drying. However, differences among response variables between years indicated that the magnitude of stream drying might be critical in intermittent streams.

Invasive species are one of the major threats to biological diversity. Invasive species of crayfish are known to negatively impact native species in aquatic ecosystems. We recently found that an Ozark endemic crayfish, Orconectes neglectus, has been introduced into the Spring River drainage of southern Missouri and northern Arkansas and appears to have the potential to negatively impact the native communities. We used quantitative kick netting along the Spring River and selected tributaries to determine the distribution and abundance of O. neglectus and its potential to impact native crayfish species. The native crayfishes Orconectes eupunctus, a species of special concern, and Cambarus hubbsi appear to no longer occur throughout much of their former range in the Spring River drainage where O. neglectus is now abundant. Orconectes eupunctus, C. hubbsi, and O. neglectus mainly used fast-flowing riffle and run habitats with a mix of gravel, cobble, and boulder, whereas the other common species collected, Orconectes punctimanus and Orconectes ozarkae, were more generalists in habitat use and were found at all sampled sites. Orconectes eupunctus and C. hubbsi were positively associated with each other, but negatively associated with O. neglectus, despite their similar habitat use. These results provide evidence that O. neglectus is expanding its range, possibly to the detriment of O. eupunctus and C. hubbsi. An intensive field survey and manipulative experiments would be required to confirm the disappearance of the native species, and the mechanisms involved.

Habitat selection by small (≤ 15.0 mm carapace length (CL)) and large (> 15.0 mm CL) individuals of the crayfish, Orconectes marchandi, O. ozarkae, O. punctimanus, and Cambarus hubbsi was examined at 4 sites in 2 streams in the Spring River watershed. Replicate quadrat samples were collected from riffle, run, pool, emergent vegetation, stream margin, and backwater habitats during the summers of 1998 and 1999. A 3-factor ANOVA was used to determine effects of habitat, species, and year on crayfish density for each site and stream. Habitat selection by crayfish was determined by comparing habitat used to habitat available for each species and size class. There were significant temporal differences in habitat used by some species-size classes of crayfish, but significant species by habitat interactions were detected at all sites for both crayfish size classes. Small O. ozarkae and O. punctimanus used mainly vegetated habitat, whereas both size classes of C. hubbsi used mainly riffles and runs. Large...

The crayfish Orconectes eupunctus, a species of special concern, appears to have been displaced from part of its former range in the Spring River drainage of Arkansas and Missouri by the invasive crayfish O. neglectus chaenodactylus. We conducted preliminary field experiments to determine whether Orconectes eupunctus were displaced from their former range and whether interspecific competition with O. neglectus chaenodactylus was responsible for the displacement. To determine whether Orconectes eupunctus were displaced from their former range by negative biotic interactions with other species we placed single adult male O. eupunctus in cages at two sites: an upstream site where O. eupunctus was formerly abundant and a downstream site where it is currently abundant. Orconectes neglectus chaenodactylus were additionally paired alongside O. eupunctus at the upstream site. We also conducted a field competition experiment with treatments consisting of (1) 3 O. eupunctus, (2) 6 O. eupunctus, and (3) 3 O. eupunctus and 3 O. neglectus chaenodactylus at a site where both species occur. In the first experiment, O. eupunctus gained significantly more weight at the downstream site than the upstream site during 2003, but not during 2002. At the upstream site, O. eupunctus showed significantly greater growth rates than O. neglectus chaenodactylus during 2002, but not during 2003. In the competition experiment, O. neglectus chaenodactylus did not significantly reduce the growth or survival of O. eupunctus. These results suggest that O. eupunctus have been displaced from their former range by biotic interactions, but interspecific competition between adult male O. eupunctus and O. neglectus chaenodactylus does not appear to be the mechanism responsible for this displacement. Further study is needed to investigate the disappearance of O. eupunctus.