Byron J. Freeman

Regional Effects of Hydrologic Alterations on Riverine Macrobiota in the New World: Tropical–Temperate Comparisons

temperate zones, paradigms and theories of how hydrologic modifications caused by dams alter the ecological dynamics of rivers are based largely on studies of temperate basins (e.g., Poff et al. 1997). Little is known about biotic responses to hydrologic modifications in tropical streams; generalizations about the effects of dams in the tropics are constrained by limited data on recently constructed, and relatively few, dams. Moreover, general ecological understanding of the effects of dams in both tropical and temperate zones is constrained by a lack of baseline information on the distribution and ecology of aquatic biota before dam construction, as well as by an overemphasis on economically important species. This article has two main objectives: to examine what is known about regional effects of hydrologic modifications in temperate and tropical areas of the New World (i.e., North and South America and the Caribbean), with an emphasis on fishes and molluscs; and to discuss research needs regarding regional effects of hydrologic alterations in temperate and tropical regions. A better understanding of regional effects of cumulative hydrologic alterations could help inform decisions on the nature and location of future hydrologic modifications. We begin with a brief description of the scope of hydrologic alterations in the New World, emphasizing dams. This is followed by a summary of biotic patterns that have emerged in hydrologically altered rivers draining temperate regions. We use the highly regulated Mobile River basin in southeastern North America as a temperate-zone case study to discuss specific biological effects. We then focus

Diversity, Distribution, and Conservation Status of the Native Freshwater Fishes of the Southern United States

Abstract The Southeastern Fishes Council Technical Advisory Committee reviewed the diversity, distribution, and status of all native freshwater and diadromous fishes across 51 major drainage units of the southern United States. The southern United States supports more native fishes than any area of comparable size on the North American continent north of Mexico, but also has a high proportion of its fishes in need of conservation action. The review included 662 native freshwater and diadromous fishes and 24 marine fishes that are significant components of freshwater ecosystems. Of this total, 560 described, freshwater fish species are documented, and 49 undescribed species are included provisionally pending formal description. Described subspecies (86) are recognized within 43 species, 6 fishes have undescribed subspecies, and 9 others are recognized as complexes of undescribed taxa. Extinct, endangered, threatened, or vulnerable status is recognized for 28% (187 taxa) of southern freshwater and diadromou...

Investigating hydrologic alteration as a mechanism of fish assemblage shifts in urbanizing streams

Abstract Stream biota in urban and suburban settings are thought to be impaired by altered hydrology; however, it is unknown what aspects of the hydrograph alter fish assemblage structure and which fishes are most vulnerable to hydrologic alterations in small streams. We quantified hydrologic variables and fish assemblages in 30 small streams and their subcatchments (area 8–20 km2) in the Etowah River Catchment (Georgia, USA). We stratified streams and their subcatchments into 3 landcover categories based on imperviousness (<10%, 10–20%, >20% of subcatchment), and then estimated the degree of hydrologic alteration based on synoptic measurements of baseflow yield. We derived hydrologic variables from stage gauges at each study site for 1 y (January 2003–2004). Increased imperviousness was positively correlated with the frequency of storm events and rates of the rising and falling limb of the hydrograph (i.e., storm “flashiness”) during most seasons. Increased duration of low flows associated with imperviousness only occurred during the autumn low-flow period, and this measure corresponded with increased richness of lentic tolerant species. Altered storm flows in summer and autumn were related to decreased richness of endemic, cosmopolitan, and sensitive fish species, and decreased abundance of lentic tolerant species. Species predicted to be sensitive to urbanization, based on specific life-history or habitat requirements, also were related to stormflow variables and % fine bed sediment in riffles. Overall, hydrologic variables explained 22 to 66% of the variation in fish assemblage richness and abundance. Linkages between hydrologic alteration and fish assemblages were potentially complicated by contrasting effects of elevated flows on sediment delivery and scour, and mediating effects of high stream gradient on sediment delivery from elevated flows. However, stormwater management practices promoting natural hydrologic regimes are likely to reduce the impacts of catchment imperviousness on stream fish assemblages.

Stream fish occurrence in response to impervious cover, historic land use, and hydrogeomorphic factors

We evaluated competing models explaining the occurrence of five stream fishes in an urbanizing watershed to determine the relative importance of (a) impervious surface and other indicators of current land use, (b) historic land use (e.g., agriculture, impoundments), and (c) hydrogeomorphic characteristics (e.g., stream size, elevation, geology). For four of five species, the best-supported models were those that included both current effective impervious cover and historic land use predictor variables, although models with only effective impervious cover were equally well supported for two of those species. For the best-supported models for three species, occurrence probability was predicted to approach zero at levels of development equivalent to about 2%-4% effective impervious cover in the surrounding region. Data were drawn from 357 fish collections made in the Etowah River basin, Georgia, USA, between 1998 and 2003 and analyzed using hi- erarchical logistic regression accounting for imperfect species detection. This is the first study we know of to examine the response of individual fish species to both increasing impervious cover and historic land use. Such individual species as- sessments will be increasingly necessary to guide policies for managing urban effects and preventing extirpations of sensi- tive species.

Application of a Multistate Model to Estimate Culvert Effects on Movement of Small Fishes

Abstract While it is widely acknowledged that culverted road-stream crossings may impede fish passage, effects of culverts on movement of nongame and small-bodied fishes have not been extensively studied and studies generally have not accounted for spatial variation in capture probabilities. We estimated probabilities for upstream and downstream movement of small (30-120 mm standard length) benthic and water column fishes across stream reaches with and without culverts at four road-stream crossings over a 4-6-week period. Movement and reach-specific capture probabilities were estimated using multistate capture-recapture models. Although none of the culverts were complete barriers to passage, only a bottomless-box culvert appeared to permit unrestricted upstream and downstream movements by benthic fishes based on model estimates of movement probabilities. At two box culverts that were perched above the water surface at base flow, observed movements were limited to water column fishes and to intervals when ...

Reproductive isolation and the expansion of an invasive hybrid swarm

Biological invasions involving hybridization proceed according to prezygotic and postzygotic reproductive isolating mechanisms. Yet few comparisons of reproductive isolation have been carried out to understand how different mechanisms prevent or promote invasions involving hybridization. Here we present a study of prezygotic and postzygotic isolation between non-native red shiner (Cyprinella lutrensis) and native blacktail shiner (C. venusta stigmatura) from the Coosa River basin (USA) to better understand the formation and expansion of invasive hybrid swarms. We conducted spawning trials to measure mating preferences and raised broods from crosses to assay hybrid viability through early juvenile development. Females of both species were more responsive to conspecific mates, although blacktail shiner females responded more often to heterospecific mates than did red shiner females. Fecundity of red shiner females was also higher than blacktail shiner females. Heterospecific crosses resulted in lower fertilization and egg hatching rates, but we found no other evidence of inviability. Rather, we found comparatively low larval mortality of F1 hybrids, which is suggestive of heterosis. These findings support prior inferences of assortative mating from genetic descriptions of hybridization, and that the invasion in the Coosa River is likely proceeding due to interspecific competition and intrinsic hybrid viability.

The Effect of Road Crossings on Fish Movements in Small Etowah Basin Streams

Abstract Increased road construction associated with urbanization may result in fragmentation and loss of fish populations in streams. In this study, we documented frequency of movement of fishes through three separate types of road-crossings (clear-span bridges, box culverts, and tube culverts) in six small streams using mark-recapture sampling. Upstream movement between areas separated by either box or tube culverts was lower than upstream movement between similar areas not separated by a road crossing. Downstream movement between areas separated by box culverts was also lower than downstream movement between areas without obstructions. Upstream and downstream movement between areas separated by clear-span bridges was generally similar to patterns of movement between areas not separated by a road crossing. Our results indicate that culverts may limit, to some degree, movements of fishes in small streams.

Accounting for Incomplete Detection when Estimating Site Occupancy of Bluenose Shiner (Pteronotropis welaka) in Southwest Georgia

Abstract Pteronotropis welaka (Bluenose Shiner) has a fragmented range throughout the Southeast, but its apparent rarity may reflect a low probability of detection during surveys. Our objectives were to obtain up-to-date status information for populations in southwest Georgia and to account for incomplete detection in our estimate of the proportion of sites occupied. We detected Bluenose Shiner at 5 of 39 sites (13%) sampled during 2004 and 2005 and estimated detection probability (p) and the proportion of sites occupied (psi) from seine-haul data. Models containing habitat covariates as predictors of p and psi provided a better description of the data than models without covariates for Bluenose Shiner and three other minnow species. Regardless of the model structure, the probability of detecting Bluenose Shiner during a single seine haul was substantially lower than for the other minnow species (3–8% vs. 13–33%). However, estimates of the proportion of sites occupied (corrected for incomplete detection) were similar to observed occupancy rates for all four species because of the large number of seine hauls we made at each site. The modeling approach we followed increased our confidence in survey results and provided information on where and how much to sample in future surveys. It has broad application to future surveys and monitoring programs for rare aquatic species in the southeastern United States.

Dealing With Uncertainty When Assessing Fish Passage Through Culvert Road Crossings

Assessing the passage of aquatic organisms through culvert road crossings has become increasingly common in efforts to restore stream habitat. Several federal and state agencies and local stakeholders have adopted assessment approaches based on literature-derived criteria for culvert impassability. However, criteria differ and are typically specific to larger-bodied fishes. In an analysis to prioritize culverts for remediation to benefit imperiled, small-bodied fishes in the Upper Coosa River system in the southeastern United States, we assessed the sensitivity of prioritization to the use of differing but plausible criteria for culvert impassability. Using measurements at 256 road crossings, we assessed culvert impassability using four alternative criteria sets represented in Bayesian belief networks. Two criteria sets scored culverts as either passable or impassable based on alternative thresholds of culvert characteristics (outlet elevation, baseflow water velocity). Two additional criteria sets incorporated uncertainty concerning ability of small-bodied fishes to pass through culverts and estimated a probability of culvert impassability. To prioritize culverts for remediation, we combined estimated culvert impassability with culvert position in the stream network relative to other barriers to compute prospective gain in connected stream habitat for the target fish species. Although four culverts ranked highly for remediation regardless of which criteria were used to assess impassability, other culverts differed widely in priority depending on criteria. Our results emphasize the value of explicitly incorporating uncertainty into criteria underlying remediation decisions. Comparing outcomes among alternative, plausible criteria may also help to identify research most needed to narrow management uncertainty.

Discordant introgression in a rapidly expanding hybrid swarm

The erosion of species boundaries can involve rapid evolutionary change. Consequently, many aspects of the process remain poorly understood, including the formation, expansion, and evolution of hybrid swarms. Biological invasions involving hybridization present exceptional opportunities to study the erosion of species boundaries because timelines of interactions and outcomes are frequently well known. Here, we examined clinal variation across codominant and maternally inherited genetic markers as well as phenotypic traits to characterize the expansion and evolution of a hybrid swarm between native Cyprinella venusta and invasive Cyprinella lutrensis minnows. Discordant introgression of phenotype, microsatellite multilocus genotype, and mtDNA haplotype indicates that the observable expansion of the C. venusta × C. lutrensis hybrid swarm is a false invasion front. Both parental and hybrid individuals closely resembling C. lutrensis are numerically dominant in the expansion wake, indicating that the non‐native parental phenotype may be selectively favored. These findings show that cryptic introgression can extend beyond the phenotypic boundaries of hybrid swarms and that hybrid swarms likely expand more rapidly than can be documented from phenotypic variation alone. Similarly, dominance of a single parental phenotype following an introduction event may lead to instances of species erosion being mistaken for species displacement without hybridization.