Search

Human activity can put non-game fishes at higher risk of extinction because of inappropriate management action. Eastern sand darter (Ammocrypta pellucida), a small benthic fish classified as threatened across much of its northern range, inhabits increasingly fragmented sandy habitats and, as a non-game fish, may be easily overlooked in conservation efforts. In this study, the authors use genotype data from nine microsatellite loci and cytochrome oxidase I (COI) sequencing data across its northern native range to re-assess genetic structure and to characterize a newly discovered, geographically disjunct population. Previous microsatellite marker analyses had identified seven distinct population genetic clusters across the region sampled; the analysis of this study showed that the newly discovered population (West Lake, Ontario) exhibits a divergent structure. COI haplotype analysis suggests that a single haplotype recolonized the Great Lakes and surrounding water bodies after the Wisconsinan glacial period, and subsequent fluctuation in water levels and habitat fragmentation resulted in divergence of genetic clusters. Although the novel West Lake population has a common ancestral source with other populations in the broader region, its divergent genetic signature merits its consideration as a separate conservation unit. The analyses of this study highlight the potential conservation implications of the discovery of new populations, particularly those of at-risk species, even within a region that has been genetically well characterized.

Translocation, the movement of organisms for conservation purposes, can result in unintended introgression if genetic material flows between populations in new ways. The Bluemask Darter Etheostoma akatulo is a federally endangered species of freshwater fish inhabiting the Caney Fork River system and three of its tributaries (Collins River, Rocky River, and Cane Creek) in Tennessee. The current conservation strategy for Bluemask Darters involves translocating the progeny of broodstock from the Collins River (in the west) to the Calfkiller River (in the east) where the species had been extirpated. In this study, we use ddRAD sequence data from across the extant range to assess this translocation strategy in light of population structure, phylogeny, and demography. We also include museum specimen data to assess morphological variation among extant and extirpated populations. Our analyses reveal substantial genetic and phenotypic disparities between a western population in the Collins River and an eastern population encompassing the Rocky River, Cane Creek, and upper Caney Fork, the two of which shared common ancestry more than 100,000 years ago. Furthermore, morphological analyses classify 12 of 13 Calfkiller River specimens with phenotypes consistent with the eastern population. These results suggest that current translocations perturb the evolutionary boundaries between two delimited populations. Instead, we suggest that repopulating the Calfkiller River using juveniles from the Rocky River could balance conflicting signatures of demography, diversity, and divergence. Beyond conservation, the microgeographic structure of Bluemask Darter populations adds another puzzle to the phylogeography of the hyperdiverse freshwater fishes in eastern North America.

Researchers studying nonmodel organisms have an increasing number of methods available for generating genomic data. However, the applicability of different methods across species, as well as the effect of reference genome choice on population genomic inference, remain difficult to predict in many cases. We evaluated the impact of data type (whole-genome vs. reduced representation) and reference genome choice on data quality and on population genomic and phylogenomic inference across several species of darters (subfamily Etheostomatinae), a highly diverse radiation of freshwater fish. We generated a high-quality reference genome and developed a hybrid RADseq/sequence capture (Rapture) protocol for the Arkansas darter (Etheostoma cragini). Rapture data from 1,900 individuals spanning four darter species showed recovery of most loci across darter species at high depth and consistent estimates of heterozygosity regardless of reference genome choice. Loci with baits spanning both sides of the restriction enzyme cut site performed especially well across species. For low-coverage whole-genome data, choice of reference genome affected read depth and inferred heterozygosity. For similar amounts of sequence data, Rapture performed better at identifying fine-scale genetic structure compared to whole-genome sequencing. Rapture loci also recovered an accurate phylogeny for the study species and demonstrated high phylogenetic informativeness across the evolutionary history of the genus Etheostoma. Low cost and high cross-species effectiveness regardless of reference genome suggest that Rapture and similar sequence capture methods may be worthwhile choices for studies of diverse species radiations.

Movement is a fundamental aspect of fish ecology, and it therefore represents an important trait to monitor for the management and conservation of fish populations. This is especially true for small benthic fish, as they often inhabit part of the catchment where their movement may be restricted by alterations to river connectivity due to human activity. Still, the movement of these small benthic fish remains poorly understood, partly because of their small size and their cryptic nature. This applies to Percilia irwini, an endangered small darter native to the south-central region of Chile. Its habitat has been affected by the presence of large hydroelectric dams and is currently threatened by the construction of several others. In this study, the authors investigated movement patterns of P. irwini from populations inhabiting different parts of the Biobío catchment, with different levels of connectivity due to natural and/or human-induced features. The authors combined chronological clustering with random forest classification to reconstruct lifelong movements from multi-elemental otolith microchemistry transects. The majority of the movements detected occurred in an undisturbed part of the catchment. These were directional upstream movements occurring between capture sites from the lower and the middle reaches of the river, representing a distance of nearly 30 km, a distance much larger than previously thought. Nonetheless, in the part of the catchment where connectivity was affected by human activity, no such movements were identified. This study shows that connectivity alteration could impede naturally occurring movement and further threaten the resilience of populations of P. irwini. Furthermore, the results presented are used to discuss advantages and disadvantages of microchemistry analysis for studying movement of small benthic fish.

Understanding a species' historical and current distribution is critical when making conservation and management decisions. Recent observations in headwater streams of northern Illinois, USA, where no previous records of Iowa Darters Etheostoma exile occurred, revealed the need to re-evaluate its state-wide distribution. We conducted a series of species distribution modelling procedures coupled with targeted field surveys to generate historical and contemporary distribution models. The historical distribution model indicated E. exile distributions were concentrated across extreme northern Illinois. Both contemporary models included results of the historical model as a model variable. Based on the initial contemporary model, 30 potential Iowa Darter sites, 10 each of three groups representing low, medium and high model scores, were sampled during the summer of 2016. Field surveys provided nine new E. exile localities and presence-absence data from field surveys were then applied to revise the contemporary model. The revised contemporary model suggests a decrease in certain areas of its historic Illinois range, as well as areas where it has potentially expanded its range.

Virginia Department of Game and Inland Fisheries; U.S. Department of Agriculture Hatch ProgramUnited States Department of Agriculture (USDA); U.S. Department of Agriculture National Institute for Food and AgricultureUnited States Department of Agriculture (USDA); U.S. Geological SurveyUnited States Geological Survey; Virginia Tech; Virginia Department of Mines, Minerals, and Energy

In the Central United States, the Arkansas darter (Etheostoma cragini) is listed as a threatened fish species by the State of Kansas. Survival of the darter is threatened by loss of habitat caused by changing streamflow conditions, in particular flow depletion. Future management of darter populations and habitats requires an understanding of streamflow conditions and how those conditions may have changed over time in response to natural and anthropogenic factors. In Kansas, streamflow alteration was assessed at 9 U.S. Geological Survey streamgages in 6 priority basins with no pronounced long-term trends in precipitation. The assessment was based on a comparison of observed (O) and predicted expected (E) reference conditions for 29 flow metrics. The O/E results indicated a likely or possible diminished flow condition in 2 basins; the primary cause of which is groundwater-level declines resulting from groundwater pumping for irrigated agriculture. In these 2 basins, habitat characteristics adversely affected by flow depletion may include stream connectivity, pools, and water temperature. The other 4 basins were minimally affected, or unaffected, by flow depletion and therefore may provide the best opportunity for preservation of darter habitat. Through the O/E analysis, anthropogenic streamflow alteration was quantified and the results will enable better-informed decisions pertaining to the future management of darters in Kansas.

Increased turbidity and the resulting decreases in water clarity have been shown to alter fish foraging behaviors in potentially detrimental ways. Given the ecological importance of darters of the genus Etheostoma and their dependence on visual cues, we examined the impact of a range of turbidity levels (3.5, 6, 10, 20, 22, and 25 nephelometric turbidity units [NTU]) on the number of prey consumed by the Tessellated Darter Etheostoma olmstedi. While a study of the Fountain Darter E. fonticola found that turbidity affected prey consumption at slightly elevated levels (≥8.7 NTU), we found no significant differences in consumption by the Tessellated Darter across turbidity levels. While there may be intragenus variation in the use of olfactory cues, another and perhaps more likely explanation is the difference in long-term exposure to a range of turbidities. The Fountain Darter is endemic to clear, primarily spring-fed rivers, whereas the Tessellated Darter is, comparatively speaking, a generalist th...

To test hypotheses explaining variation in elaborate male colouration across closely related species groups, ancestral-state reconstructions and tests of phylogenetic signal and correlated evolution were used to examine the evolution of male body and fin colouration in a group of sexually dichromatic stream fishes known as darters (Percidae: Etheostomatinae). The presence or absence of red-orange and blue-green male colour traits were scored across six body regions in 99 darter species using a recently estimated amplified fragment length polymorphism (AFLP) phylogeny for comparative analyses. Ancestral-state reconstructions infer the most recent common ancestor of darters to lack red-orange colour and possess blue-green colour on different body regions, suggesting variation between species is due to independent gains of red-orange and losses of blue-green. Colour traits exhibit substantial phylogenetic signal and are highly correlated across body regions. Comparative analyses were repeated using an alternative phylogenetic hypothesis based on one mitochondrial and two nuclear genes, yielding similar results to analyses based on the AFLP phylogeny. Red-orange colouration in darters appears to be derived; whereas, blue-green appears to be ancestral, which suggests that different selection mechanisms may be acting on these two colour classes in darters.

Understanding how biotic and abiotic processes influence community assembly is a fundamental aim in ecology. Although spatial scales at which communities are studied may affect the relative importance of such assembly processes, spatial influences on community assembly have not been thoroughly addressed. We tested how spatial scale affects inferences of habitat filtering and competitive exclusion assembly processes in darter (Percidae: Etheostomatinae) assemblages across four temperate stream systems. We predicted competitive exclusion would influence assembly in fine-scale assemblages, and habitat filtering would be more influential as spatial scale increased. Moreover, we assumed that habitat heterogeneity would increase with scale, and consequently alleviate direct competitive exclusion acting at finer scales. Using a framework that incorporated genetic relatedness, morphological traits, and habitat use among co-occurring darter species, we identified ecological and evolutionary patterns of structure, which allowed us to elucidate processes of assembly. Based on phylogenetic structure, assemblages showed an increase in habitat filtering (i.e., increased phylogenetic clustering) as we scaled up from fine to intermediate assemblages; however, we found mixed signals for habitat filtering and competitive exclusion at the broadest spatial scale. While habitat filtering was found to have an overall high relative importance during assembly, we also found influence of competitive exclusion processes based on limited morphological similarity among co-occurring species. Our results generally support an increased influence of habitat filtering processes in broader scale assemblages. Moreover, we suggest that habitat filtering and competitive exclusion processes act simultaneously during assembly, although the relative influence of each process may be spatial-scale dependent.

Sixteen darter species, including the federally endangered Boulder Darter Etheostoma wapiti, are known to occur in the Elk River, a large, flow-regulated tributary of the Tennessee River, Tennessee–Alabama. Since the construction of Tims Ford Dam (TFD) in 1970, habitat modification caused by cold, hypolimnetic water releases and peak-demand hydropower generation has contributed to population declines and range reductions for numerous aquatic species in the main-stem Elk River. We developed Bayesian hierarchical multispecies occupancy models to determine the influence of site- and species-level characteristics on darter occurrence by using presence–absence data for 15 species collected from 39 study sites. Modeling results indicated that large-river obligate species, such as the Boulder Darter, were 6.92 times more likely to occur for every 37-km increase in the distance downstream from TFD. In contrast, small-stream species were 2.35 times less likely and cosmopolitan species were 1.88 times less ...

Elaborate, sexually dimorphic traits are widely thought to evolve under sexual selection through female preference, male-male competition, or both. The orangethroat darter (Etheostoma spectabile) is a sexually dichromatic fish in which females exhibit no preferences for male size or coloration. We tested whether these traits affect individual reproductive success in E. spectabile when multiple males are allowed to freely compete for a female. The quality and quantity of male coloration were associated with greater success in maintaining access to the female and in spawning as the primary male (first male to participate). On the other hand, sneaking behavior showed little correlation with coloration. Male breeding coloration in E. spectabile may therefore demonstrate how intrasexual competition can be a predominant factor underlying the evolution of male ornaments.

Closely related animal lineages often vary in male coloration, and ecological selection is hypothesized to shape this variation. The role of ecological selection in inhibiting male color has been documented extensively at the population level, but relatively few studies have investigated the evolution of male coloration across a clade of closely related species. Darters are a diverse group of fishes that vary in the presence of elaborate male nuptial coloration, with some species exhibiting vivid color patterns and others mostly or entirely achromatic. We used phylogenetic logistic regression to test for correlations between the presence/absence of color traits across darter species and the ecological conditions in which these species occur. Environmental variables were correlated with the presence of nuptial color in darters with colorful species tending to inhabit environments that would support fewer predators and potentially transmit a broader spectrum of natural light compared to species lacking male coloration. We also tested the color preferences of a common darter predator, largemouth bass, and found that it exhibits a strong preference for red, providing further evidence of predation as a source of selection on color evolution in darters. Ecological selection therefore appears to be an important factor in dictating the presence or absence of male coloration in this group of fishes.

Habitat partitioning among syntopic darter species (Percidae: Etheostomatini) was examined in the Flint River and Estill Fork of the Paint Rock River in northeast Alabama from September 2010 to September 2011. Substrate composition, depth, and water velocity were important variables by which species segregated. Niche overlap values also indicated a high degree of habitat partitioning among species at both study sites, although a substantial amount of overlap was seen between Etheostoma simoterum and Etheostoma caeruleum at the Estill Fork site. Segregation among species was present throughout the year, but the significance of differences in habitat preferences did vary seasonally, especially during the post-spawn period when habitat heterogeneity was reduced. Intra-specific variation in habitat utilization was present among some species, with males generally preferring higher flow and coarser substrates than females. Habitat segregation was a well-structured mechanism allowing the coexistence of ecologically similar species. Copyright © 2015 John Wiley & Sons, Ltd.

Environmental variation can shape phenotypic variation in organisms, but most evidence for trait differentiation comes from analyses of dichotomous habitat types that differ in only one or few key environmental factors. In reality, environmental variation is often more subtle, gradual and multifarious. Here, we investigated geographic variation in body shape of two darter species (Etheostoma spectabile and Etheostoma flabellare; Percidae) that occur along river gradients. This study addressed three specific questions: Is there intraspecific geographic variation in the two species across different sites in the Ozark Highlands of Oklahoma (USA)? Is phenotypic variation across sites correlated with abiotic environmental conditions? Do the two species share site-specific (i.e. convergent) phenotypic variation in areas where they occur together? Our results indicated significant body shape variation in both species. Population differences in body shape were particularly correlated with variation in substrate composition. The combined analysis of both species indicated a small but significant effect of convergence on body shape wherever they are sympatric; shared variation, however, was not related to any environmental variables included in the analysis. While it remains unclear whether phenotypic variation in these species is due to heritable differentiation or environmentally induced plasticity, our results indicate that even subtle and gradual environmental variation can induce substantial variation in phenotypes on a relatively small spatial scale.

It is not known how environmental pressures and sexual selection interact to influence the evolution of extravagant male traits. Sexual and natural selection are often viewed as antagonistic forces shaping the evolution of visual signals, where conspicuousness is favored by sexual selection and crypsis is favored by natural selection. Although typically investigated independently, the interaction between natural and sexual selection remains poorly understood. Here, we investigate whether sexual dichromatism evolves stochastically, independent from, or in concert with habitat use in darters, a species-rich lineage of North American freshwater fish. We find the evolution of sexual dichromatism is coupled to habitat use in darter species. Comparative analyses reveal that mid-water darter lineages exhibit a narrow distribution of dichromatism trait space surrounding a low optimum, suggesting a constraint imposed on the evolution of dichromatism, potentially through predator-mediated selection. Alternatively, the transition to benthic habitats coincides with greater variability in the levels of dichromatism that surround a higher optimum, likely due to relaxation of the predator-mediated selection and heterogeneous microhabitat dependent selection regimes. These results suggest a complex interaction of sexual selection with potentially two mechanisms of natural selection, predation and sensory drive, that influence the evolution of diverse male nuptial coloration in darters.

Quantifying spatial genetic structure can reveal the relative influences of contemporary and historic factors underlying localized and regional patterns of genetic diversity and gene flow – important considerations for the development of effective conservation efforts. Using 10 polymorphic microsatellite loci, we characterize genetic variation among populations across the range of the Eastern Sand Darter (Ammocrypta pellucida), a small riverine percid that is highly dependent on sandy substrate microhabitats. We tested for fine scale, regional, and historic patterns of genetic structure. As expected, significant differentiation was detected among rivers within drainages and among drainages. At finer scales, an unexpected lack of within-river genetic structure among fragmented sandy microhabitats suggests that stratified dispersal resulting from unstable sand bar habitat degradation (natural and anthropogenic) may preclude substantial genetic differentiation within rivers. Among-drainage genetic structure indicates that postglacial (14 kya) drainage connectivity continues to influence contemporary genetic structure among Eastern Sand Darter populations in southern Ontario. These results provide an unexpected contrast to other benthic riverine fish in the Great Lakes drainage and suggest that habitat-specific fishes, such as the Eastern Sand Darter, can evolve dispersal strategies that overcome fragmented and temporally unstable habitats.

Understanding linkages among fluvial geomorphology, habitat, and aquatic biota is critical for effective stream ecosystem conservation. However, composite effects of hydrogeomorphic adjustment and condition, which collectively represent channel stability, on freshwater mussel and stream fish assemblages remain unresolved. Associations between stream hydrogeomorphic characteristics (channel geometry, substrate composition, stream flow) and mussel and stream fish assemblages were explored at 20 study reaches characterized by riffle–pool interfaces (RPIs) in Ohio, USA. At a coarse resolution using categorical classifications of equilibrium (i.e. stable) vs. adjusting (i.e. unstable) RPIs, overall fish and darter density was greater at adjusting RPIs (P = 0.048 and P = 0.024, respectively). Conversely, fish species richness was 1.2× greater at equilibrium than adjusting RPIs (P = 0.047). Analysis of quantitative hydrogeomorphic data collected with fine-resolution surveys showed that hydrogeomorphic parameters explained from 20% (darter assemblage evenness) to 55% (density of mussels known to use darters as hosts) of the variation observed in all assemblages. Drainage area was significant in most models with variable influence: R2 = 0.10 for darter species richness to R2 = 0.41 for Simpson's diversity index of mussels with darter hosts. Other important predictor variables included embeddedness, velocity, shear stress, roughness, channel dimensions, and sediment size. Whereas coarse-level fluvial geomorphic classifications may be meaningful for fish, they appear less so for mussels. Fine-resolution quantitative hydrogeomorphic variables provided substantially more information for both assemblages, although hydrogeomorphology–fish and hydrogeomorphology–mussel relationships were not consistent. Some of the strongest relationships related to mussels that use darters as hosts, suggesting that these species are particularly sensitive to hydrogeomorphic conditions. Collectively, these results indicate that fluvial geomorphic condition and characteristics can simultaneously influence co-dependent stream biota. Stream conservation and management plans that include explicit hydrogeomorphic surveys may appreciably benefit cohabitating freshwater fish and mussel assemblages. Copyright © 2014 John Wiley & Sons, Ltd.

The dispersal ecology of most stream fishes is poorly characterised, complicating conservation efforts for these species. We used microsatellite DNA marker data to characterise dispersal patterns and effective population size (Ne) for a population of Roanoke logperch Percina rex, an endangered darter (Percidae). Juveniles and candidate parents were sampled for 2 years at sites throughout the Roanoke River watershed. Dispersal was inferred via genetic assignment tests (ATs), pedigree reconstruction (PR) and estimation of lifetime dispersal distance under a genetic isolation-by-distance model. Estimates of Ne varied from 105 to 1218 individuals, depending on the estimation method. Based on PR, polygamy was frequent in parents of both sexes, with individuals spawning with an average of 2.4 mates. The sample contained 61 half-sibling pairs, but only one parent–offspring pair and no full-sib pairs, which limited our ability to discriminate natal dispersal of juveniles from breeding dispersal of their parents between spawning events. Nonetheless, all methods indicated extensive dispersal. The AT indicated unrestricted dispersal among sites ≤15 km apart, while siblings inferred by the PR were captured an average of 14 km and up to 55 km apart. Model-based estimates of median lifetime dispersal distance (6–24 km, depending on assumptions) bracketed AT and PR estimates, indicating that widely dispersed individuals do, on average, contribute to gene flow. Extensive dispersal of P. rex suggests that darters and other small benthic stream fishes may be unexpectedly mobile. Monitoring and management activities for such populations should encompass entire watersheds to fully capture population dynamics.

During the past 100 years, most large rivers in North America have been altered for flood control, hydropower, navigation or water supply development. Although these activities clearly provide important human services, their associated environmental disturbances can profoundly affect stream-dwelling organisms. We used dynamic multi-species occupancy models combined with a trait-based approach to estimate the influence of site-level and species-level characteristics on patch dynamic rates for 15 darter species native to the Elk River, a large, flow-regulated Tennessee River tributary in Tennessee and Alabama. Dynamic occupancy modelling results indicated that for every 2.5 °C increase in stream temperature, darters were 3.94 times more likely to colonize previously unoccupied stream reaches. Additionally, large-bodied darter species were 3.72 times more likely to colonize stream reaches compared with small-bodied species, but crevice-spawning darter species were 5.24 times less likely to colonize previously unoccupied stream reaches. In contrast, darters were 2.21 times less likely to become locally extinct for every 2.5 °C increase in stream temperature, but high stream discharge conditions elevated the risk of local extinction. Lastly, the presence of populations in neighbouring upstream study reaches contributed to a lower risk of extinction, whereas the presence of populations in neighbouring downstream study reaches contributed to higher rates of colonization. Our study demonstrates the application of a trait-based approach combined with a metapopulation framework to assess the patch dynamics of darters in a regulated river. Results from our study will provide a baseline for evaluating the ecological consequences of alternative dam operations. Copyright © 2014 John Wiley & Sons, Ltd.

Robust reproductive isolation was found between the rainbow darter Etheostoma caeruleum and the orangethroat darter Etheostoma spectabile, as more offspring were produced when conspecific males and females were crossed as compared with heterospecific crosses. Furthermore, fewer eggs resulted from heterospecific crosses involving sympatric E. spectabile females than those using allopatric E. spectabile females, while a similar pattern was not observed in heterospecific crosses using E. caeruleum females. These results suggest that reinforcement, i.e. selection for pre-zygotic reproductive barriers driven by reduced hybrid fitness, may have contributed to the evolution and maintenance of reproductive barriers between these potentially hybridizing species in sympatry.

This study examined how colour varies across season and sex in the fantail darter Etheostoma flabellare and the banded darter Etheostoma zonale. Etheostoma flabellare has male-only parental care and exhibited slight sexual dimorphism in overall colour, with no discernible effect of season on colour; whereas E. zonale does not have parental care and exhibited substantial sexual dimorphism in colour, but only in the breeding season. Additionally, antipredator behaviour of E. zonale was compared between males that were fully coloured during the breeding season and males that were partially coloured at that time, but the effects of colour and season were not consistent across males.

In this study, female preferences and behavioural isolation were estimated in a pair of allopatric sister species, Etheostoma duryi and Etheostoma flavum. Dichotomous mate preference trials were conducted to determine whether females prefer to associate with conspecific over heterospecific males and free-spawning assays were conducted to determine whether those preferences translated into behavioural isolation. Dichotomous mate choice trials revealed asymmetric female preference, as female E. flavum preferred conspecific males, whereas female E. duryi showed no preference. Free-spawning assays indicated that behavioural isolation remains incomplete between E. duryi and E. flavum (IB = 0·19). In addition to female mating preferences, male behaviour also appeared to influence mating outcomes as male E. flavum consistently courted conspecific females more often in free-spawning assays whereas male E. duryi did not. The data therefore suggest that despite marked divergence in male nuptial colour, divergence in female preferences between these species may not be sufficient to maintain species boundaries upon secondary contact. These results contrast with similar work in a sympatric darter species pair and may be explained by considering the contributions of reinforcement and differences in colour pattern as well as colour value.

Connectivity throughout large riverine networks is often compromised by lock and dam (L/D) structures designed to facilitate year-round navigation. The resultant interruption of flow inhibits free passage of aquatic biota potentially isolating mainstem and tributary communities. Our objectives were (i) to evaluate the impact of a series of navigational L/D structures on two targeted fish assemblages (TFAs): large-bodied (>250 mm total length) pelagic and benthic (darter) communities and (ii) to examine patterns of tributary and mainstem connectivity. We systematically captured fishes utilizing gillnets, benthic trawls and backpack electrofishers from an impounded and a free-flowing reach extending over 203 km of the Allegheny River in Pennsylvania. Species richness among the large-bodied targeted fish assemblage was distributed somewhat evenly throughout each pool, peaking near dams and in an undredged pool. Depauperate darter assemblages of low species richness characterized most of the navigable reach with diversity and abundance concentrated in L/D tailrace areas. By contrast, darter communities inhabiting the free-flowing reach were more diverse, abundant and evenly distributed, indicating the influence of uninterrupted connectivity. Community similarity at mainstem/tributary junctions increased with increasing tributary size with 50% of the mainstem species complement also inhabiting the largest tributary. This study underscores the importance of dams as barriers to ichthyofaunal connectivity, particularly to those benthic fishes which are relatively immobile and habitat specific, and the importance of mainstem/tributary junctions as avenues of riverine connectivity. Copyright © 2013 John Wiley & Sons, Ltd.

The only known extant population of the diamond darter (Crystallaria cincotta) exists in the lower 37 km of Elk River, WV, USA. Our understanding of diamond darter habitat use was previously limited, because few individuals have been observed during sampling with conventional gears. We quantified microhabitat use of diamond darters based on measurements of water depth, water velocity and per cent substrate composition. Using spotlights at night-time, we sampled 16 sites within the lower 133 km of Elk River and observed a total of 82 diamond darters at 10 of 11 sampling sites within the lower 37 km. Glides, located immediately upstream of riffles, were the primary habitats sampled for diamond darters, which included relatively shallow depths (

Aim Rare aquatic species are a substantial component of biodiversity, and their conservation is a major objective of many management plans. However, they are difficult to assess, and their optimal habitats are often poorly known. Methods to effectively predict the likely locations of suitable rare aquatic species habitats are needed. We combine two modelling approaches to predict occurrence and general abundance of several rare fish species. Location Allegheny watershed of western New York State (USA) Methods Our method used two empirical neural network modelling approaches (species specific and assemblage based) to predict stream-by-stream occurrence and general abundance of rare darters, based on broad-scale habitat conditions. Species-specific models were developed for longhead darter (Percina macrocephala), spotted darter (Etheostoma maculatum) and variegate darter (Etheostoma variatum) in the Allegheny drainage. An additional model predicted the type of rare darter-containing assemblage expected in each stream reach. Predictions from both models were then combined inclusively and exclusively and compared with additional independent data. Results Example rare darter predictions demonstrate the method's effectiveness. Models performed well (R2 ≥ 0.79), identified where suitable darter habitat was most likely to occur, and predictions matched well to those of collection sites. Additional independent data showed that the most conservative (exclusive) model slightly underestimated the distributions of these rare darters or predictions were displaced by one stream reach, suggesting that new darter habitat types were detected in the later collections. Main conclusions Broad-scale habitat variables can be used to effectively identify rare species' habitats. Combining species-specific and assemblage-based models enhances our ability to make use of the sparse data on rare species and to identify habitat units most likely and least likely to support those species. This hybrid approach may assist managers with the prioritization of habitats to be examined or conserved for rare species.

To inquire how male size interacts with alloparental behaviour and mating success in the tessellated darter Etheostoma olmstedi, males were given a choice of nests with or without eggs; subsequent nest occupancy, takeovers and egg deposits were monitored. Subordinate males readily occupied available nests with eggs but were often evicted by dominant males, suggesting that males of all sizes compete for the opportunity to provide allopaternal care in this species.

Five darter species (Greenside Darter Etheostoma blennioides, Rainbow Darter E. caeruleum, Fantail Darter E. flabellare, Johnny Darter E. nigrum, and Orangethroat Darter E. spectabile) were collected from seven streams and rivers in two Indiana watersheds to determine patterns of microhabitat selection with respect to depth, water velocity (flow), and substrate size. Heterogeneous habitats create a variety of niche dimensions, promoting the coexistence of congeneric species. Greenside and Rainbow darters were most commonly found among intermediate substrate sizes (cobble + boulder) and in locations with higher velocities and deeper water. Fantail and Orangethroat darters associated with intermediate to large substrate sizes (cobble + bedrock) were found in shallower water with lower velocities. Fantail Darters were only observed in the Whitewater River watershed. In contrast, Johnny Darters were observed only within the White River watershed among small substrate sizes (silt + sand), moderate vel...

– The ashy darter, Etheostoma cinereum, is an imperiled fish within the Cumberland and Tennessee drainages of the southeast United States. An understanding of habitat associations and the relationship of habitat use across multiple spatial scales are critical elements in its conservation. Our objectives were to quantify habitat associations at the stream reach and microhabitat scales for adult and juvenile darters, and to understand the linkage between the two scales based on gradients of habitat use. We focused our efforts within the Rockcastle River, Kentucky (Cumberland River drainage), because the watershed was known to contain a relatively large ashy darter population. Three hundred twenty-two individuals were collected from 21 reaches. The species was restricted to the mainstem of the river and the lower reaches of the larger tributaries. The distribution and abundance of adult and juvenile darters differed significantly at each spatial scale, and both groups demonstrated non-random use of the available habitat. Gradients of stream size and substrate size were identified as important factors. A threshold of environmental quality was determined based on the habitat use patterns among the two scales. Habitat use between the two scales was independent within the threshold, indicating that the specific quality of the microhabitats did not necessarily matter within a stream reach. However, beyond the threshold, a decrease of at least 48% in adult and juvenile darter abundance was seen, indicating that a sufficient network of suitable microhabitats is needed to support a good population of darters within a stream reach.

Eastern sand darter (Ammocrypta pellucida) is listed as Threatened under the Canadian Species at Risk Act. Little relevant biological information is available for most Canadian populations and only limited information is available for populations in the United States. To supplement the paucity of information, this study collected biological information on eastern sand darter during field surveys in 2006-2007 from 10 sites in the lower Thames River, Ontario, Canada. Collected data were used to estimate critical life history traits including: longevity, fecundity, clutch size and number, growth, survival, age-at-first-maturity and cohort age structure. Longevity was 3+ years, with age-at-first-maturity being 1+ for both sexes. Examination of egg sizes from preserved females in 2007 showed a bimodal distribution, suggesting two spawning occurrences of 71 eggs each. Quantitative comparison of lower Thames River biological information with a more southerly eastern sand darter population in the Little Muskingum River, Ohio, demonstrated little variation between the populations. Data comparisons suggest that localised environmental factors are affecting biological traits (growth), in particular water temperature that may be controlled by differences in riparian cover and/or groundwater input.

Introgressive hybridization and incomplete lineage sorting complicate the inference of phylogeny, and available species-tree methods do not simultaneously account for these processes. Both hybridization and ancestral polymorphism have been invoked to explain divergent phylogenies inferred from different datasets for Stigmacerca, a clade of 11 North American darter species. Species of Stigmacerca are characterized by a mating system involving parental care with males guarding nesting territories and fertilized eggs. Males of four species of Stigmacerca develop egg-mimic nuptial structures on their second dorsal fins during the breeding season. Previous phylogenies suggest contrasting scenarios for the evolution of this nuptial trait. Using a combination of coalescent-based methods, we analyzed a dataset comprising a mitochondrial gene and 15 nuclear loci to estimate relationships and simultaneously test for introgressive hybridization. Our analyses identified several instances of interspecific gene flow involving both cytoplamsmic haplotypes and nuclear alleles. The new phylogeny was used to infer a single origin and recent loss of egg-mimic structures in Stigmacerca and led to the discovery of a phylogenetically distinct species. Our results highlight the limited strategies available to account for introgressive hybridization in the inference of species relationships and the likely effects of this process on reconstructing trait evolution.

Like many fishes native to western Great Plains streams, the Arkansas darter Etheostoma cragini has declined, apparently in response to changes in flow regimes and habitat fragmentation. We investigated the effectiveness of translocation as a management strategy to conserve this threatened species in the Arkansas River basin of southeastern Colorado. We used a multiscale design to sample the darter and several attributes of its habitat at the local 10-m site scale, the 3.25-km translocation segment scale, and the 10-km riverscape scale in all 19 streams where it had been previously translocated. The darter was captured in 11 of the 19 streams, although 5 were completely dry when visited. Arkansas darters had reproduced in 10 of the 11 streams (one criterion in the state recovery plan), and 6 streams also met a second criterion for abundance (>500 individuals). However, the populations in only two streams unequivocally met a third criterion of being self-sustaining, because the other four streams ...

Male-only care is the most frequent parental care behaviour in teleost fishes, but little is known about its evolutionary origins and patterns of diversity in species-rich lineages. Darters are a clade of North American freshwater fishes that contain both nonparental care species and species with male-only care. In darters, paternal care takes the form of egg-guarding and other egg-tending behaviours that are dependent on the female mode of egg deposition. Male care has been hypothesized to evolve independently in darters at least three times, and it has been thought to be irreversible. We investigated the diversification of egg-deposition behaviours and the evolution of complex male care using published descriptions of darter reproductive behaviours and a multilocus molecular phylogeny that included all 146 species for which reproductive behaviours are known. We find support for two origins of male-only care behaviour. One origin of paternal care occurred relatively early in the radiation of Etheostoma and is characteristic of a recently discovered clade, Goneaperca. The other origin of male-only care occurred much more recently in a derived clade of Nothonotus. Our analyses of character diversification demonstrate reversals from care to noncare and multiple transitions between egg-deposition behaviours that are not associated with parental care.

Rivers and streams are among the most threatened ecosystems worldwide, and their fish assemblages have been modified by anthropogenic habitat alteration and introductions of non-native species. Consequently, two frequently observed patterns of assemblage change over time are species loss and biotic homogenization. In the present study, we compared contemporary (2006–2007) and historical (1948–1955) assemblages of darters, a group of small benthic fishes of the family Percidae, in the Arkansas River drainage of northeastern Oklahoma, USA. Results showed species loss between the two sampling periods, with historical estimates of overall species diversity across the study area exceeding contemporary estimates by five to eight species. Assemblages showed a low degree of darter similarity based on species presence and absence, with pairwise site comparisons (Jaccard’s similarity index) between historical and contemporary samples averaging  0.35. No significant homogenization or differentiation of assemblages occurred. Range expansion of widespread species, one of the primary mechanisms of biotic homogenization, was not observed; rather, all species occurred at a smaller proportion of sites in contemporary samples. Our results highlight the threat posed by anthropogenic habitat alteration to taxonomic groups such as darters, most of which are habitat specialists. However, our results suggest that biotic homogenization is unlikely to occur in the absence of immigration, especially if assemblages are subjected to ‘novel disturbances’ such as dam construction and watershed-scale habitat degradation which negatively affect all components of the assemblage.

Adaptive cluster sampling (ACS) designs were tested against simple random sampling (SRS) designs to determine whether ACS designs were more suitable sampling protocols for monitoring rare fish species. To test the utility of these designs, baseline data on the tuxedo darter Etheostoma lemniscatum (a rare, federally endangered fish) were collected at three sites on the Big South Fork of the Cumberland River and used in computer simulation of ACS designs. Based on the simulation models, five ACS designs were chosen and tested at 13 potential monitoring sites. In terms of efficiency, ACS designs performed better than SRS designs, providing estimates with smaller standard errors. Adaptive cluster sampling design efficiency increased with effort; therefore, the two goals—minimizing effort and maximizing accuracy—are incompatible. Because it had less error, an inverse ACS design was recommended, although it required more sampling effort than other designs. Factors possibly affecting overall design perf...

We characterised microhabitat availability and use by adult and young-of-year (YOY) snail darters (Percina tanasi Etnier 1976) while snorkelling in the French Broad and Hiwassee rivers, TN, USA. Both age groups of snail darters disproportionately used most microhabitat variables compared to their availability. Snail darters primarily occupied moderately deep, swift water over gravel substrates with little macrophyte coverage and no silt. Univariate comparisons indicated that adult and YOY darters occupied different habitat, but there was no marked differences between principal components analysis plots of multivariate microhabitat use within a river. Although the availability of microhabitat variables differed between the French Broad and Hiwassee rivers, univariate means and multivariate plots illustrated that the habitats used were generally similar by age groups of snail darters between rivers. Because our observations of habitat availability and use were constrained to low flow periods and depths

Hypotheses of diversification in eastern North American freshwater fishes have focused primarily on allopatric distributions of species between disjunct highland areas and major river systems. However, these hypotheses do not fully explain the rich diversity of species within highland regions and river systems. Relatively old diversification events at small geographic scales have been observed in the Barcheek Darter subclade that occurs in the Cumberland River drainage (CRD) in Kentucky and Tennessee, United States of America, but it is unknown if this pattern is consistent in other darter subclades. We explored phylogeographic diversity in two species of Nothonotus darters, N. microlepidus and N. sanguifluus, endemic to the CRD to compare phylogenetic patterns between Barcheek Darters and species of Nothonotus. We collected sequence data for a mitochondrial gene (cytb) and three nuclear genes (MLL, S7 and RAG1) from 19 N. microlepidus and 35 N. sanguifluus specimens. Gene trees were estimated using maximum likelihood and Bayesian methods, and a 'species tree' was inferred using a Bayesian method. These trees indicate that species diversity in Nothonotus is underestimated. Five distinct lineages were evident, despite retained ancestral polymorphism and unsampled extirpated populations. Comparison of chronograms for Barcheek Darters and Nothonotus revealed that microendemism resulting from species diversification at small geographic scales in the CRD is a consistent pattern in both old and young darter subclades. Our analyses reveal that geographic isolating mechanisms that result in similar phylogeographic patterns in the CRD are persistent through long expanses of evolutionary time.

Elaborate visual communication signals characterize many animal lineages. Often sex-limited, these signals are generally assumed to result from sexual selection, and in many cases, their evolution is thought to play a central role in speciation. The co-evolution of male visual signals and female preferences is hypothesized to result in behavioral isolation between divergent lineages; however, for many lineages characterized by elaborate visual signals, the importance of visual differences in behavioral isolation is not well established. Darters (fish genus Etheostoma) are particularly appropriate for examining the role of visual signals in behavioral isolation. They comprise one of the most diverse groups of North American freshwater fish, and nearly every species is characterized by unique nuptial coloration. Multiple darter species co-exist in sympatric populations, indicating that reproductive barriers are central to maintaining these extraordinarily diverse color patterns. This study demonstrates the presence of behavioral isolation between a pair of distinctly colored sympatric darter species, Etheostoma barrenense and Etheostoma zonale, through experimental observations using an artificial stream. In addition, a series of dichotomous mate-choice trials indicate that females prefer conspecific males over heterospecifics based on visual differences alone. We therefore provide the first evidence that visual signals are a critical trait maintaining behavioral isolation in darters, a lineage of fishes with spectacular diversification in visual communication.

Using mark-recapture methods, we studied movements of endangered Roanoke logperch (Percina rex Jordan & Evermann), a benthic darter, at 12 riffle-run sites over a 9-year-long period in the Roanoke River, Virginia, USA. Our primary objective was to characterise movements among transects within sites, but we opportunistically recorded two between-site movements as well. Our recapture rate was low (22 of 485 marked fish), but most recaptured fish exhibited movements between tagging and recapture, relocating either to another transect within a site (12 fish) or to another site altogether (two fish). Within sites, Roanoke logperch exhibited fidelity over time to the areas in which they were initially marked. These restricted areas were lengthier than transects (>15 m) but shorter than entire riffle-runs (

In order to investigate a potential hybrid zone between the candy darter, Etheostoma osburni, and variegate darter, Etheostoma variatum, and examine population variation within E. osburni, a suite of primers for 15 polymorphic microsatellite loci were developed. The average number of alleles per locus was 5.5 in E. osburni and 7.6 in E. variatum, and the average observed heterozygosities were 62.5% and 71.4%, respectively. There were no deviations from Hardy-Weinberg equilibrium and no observed linkage disequilibrium after Bonferroni correction. The utility of these primers was also tested in 11 species of darters representing all four genera of darters. Success of cross-species amplification was largely consistent with phylogenetic relationships of darters.

The Roanoke logperch (Percina rex Jordan and Evermann), an endangered fish, occurs in only six watersheds in the Roanoke and Chowan river drainages of Virginia, USA. The species' population genetic structure is poorly known. We developed 16 microsatellite markers that were reliably scorable and polymorphic P. rex. Markers were also screened in seven other darter species of the genus Percina. Most markers exhibited successful amplification and polymorphism in several species. These markers may therefore prove useful for population genetic studies in other darters, a diverse but highly imperiled group.

The effects of changing buoyancy on the diving and feeding behaviour of Reed Cormorants Phalacrocorax africanus and Darters Anhinga melanogaster was investigated at Lake Kariba, Zimbabwe. A qualitative model of the energetic constraints caused by buoyancy changes on the diving behaviour of these two birds is presented and the predictions from the model are tested. The buoyancy of both species declined exponentially at different rates with water depth. Reed Cormorants were neutrally buoyant at 5–6 m while Darters were neutrally buoyant at 2–4 m depth. Buoyancy changes affect underwater swimming speed, which for Reed Cormorants is twice as fast on the bottom than when commuting, and for the Darter is significantly slower when diving than at any other time. Cormorants feeding in water deeper then 6 m spent less time on the bottom and fed less successfully than those birds feeding in shallower water. This is because their bottom times were significantly reduced as a result of the energetic constraints caused by changes in their buoyancy.

– We compared gonadosomatic index (GSI) and histological analysis of ovaries for identifying reproductive periods of fishes to determine the validity of using GSI in future studies. Four small-bodied riverine species were examined in our comparison of the two methods. Mean GSI was significantly different between all histological stages for suckermouth minnow and red shiner. Mean GSI was significantly different between most stages for slenderhead darter; whereas stages 3 and 6 were not significantly different, the time period when these stages are present would allow fisheries biologists to distinguish between the two stages. Mean GSI was not significantly different for many histological stages in stonecat. Difficulties in distinguishing between histological stages and GSI associated with stonecat illustrate potential problems obtaining appropriate sample sizes from species that move to alternative habitats to spawn. We suggest that GSI would be a useful tool in identifying mature ovaries in many small-bodied, multiple-spawning fishes. This information could be combined with data from histology during mature periods to pinpoint specific spawning events.

Incongruence between recognized taxonomy and phylogenetic relationships between two species from a diverse clade (Percidae: Etheostomatinae) of stream fishes was found in a mitochondrial (mt) DNA gene tree. Two darters in subgenus Oligocephalus, Etheostoma uniporum current darter and Etheostoma caeruleum rainbow darter were sampled throughout their sympatric distribution in the Ozark Highlands of the central United States. Sequences from cytochrome (cyt) b and the first intron of the nuclear marker S7 were analysed separately using maximum parsimony and Bayesian methods. Cyt b recovered both species as polyphyletic; E. uniporum haplotypes were interspersed within E. caeruleum. However, both species were monophyletic and non-sister taxa based on S7. The cyt b gene tree pattern is caused by introgressive hybridization resulting in the mtDNA replacement of E. uniporum haplotypes by those of E. caeruleum. Some E. uniporum haplotypes are shared with geographically proximate E. caeruleum, and this is consistent with recent hybridization, while other E. uniporum haplotypes indicate historical sorting of introgressed lineages. The mechanism of introgression is likely asymmetric sneaking behaviour by male E. uniporum, a mating tactic observed in related species. MtDNA replacement may have occurred in E. uniporum due to drift fixation in a historically small female effective population. Additional evidence for darter hybridization will likely be discovered in future molecular genetic surveys of the nearly 200 species in eastern North America.

Identifying the manner in which reproductive barriers accumulate during lineage divergence is central to establishing general principles of species formation. One outstanding question is which isolating mechanisms form the first complete barrier to gene flow in a given lineage or under a particular set of conditions. To identify these initial reproductive barriers requires examining lineages in very early stages of divergence, before multiple reproductive barriers have evolved to completion. We quantified the strength of three postmating barriers in a pair of darter species and compared these estimates to each other and to the strength of behavioral isolation (BI) reported in a previous study. Results reveal no evidence of gametic incompatibility but intermediate levels of conspecific sperm precedence and hybrid inviability. As BI is nearly complete, our analysis comparing the strength of multiple reproductive barriers implicates the evolution of mate choice as central to both the origin and maintenance of these species. Further examination of ecological isolation and hybrid sterility is necessary to determine the role of these barriers in darter speciation.

Diving birds have to overcome buoyancy, especially when diving in shallow water. Darters and anhingas (Anhingidae) are specialist shallow-water divers, with adaptations for reducing their buoyancy. Compared to closely-related cormorants (Phalacrocoracidae), darters have fully wettable plumage, smaller air sacs and denser bones. A previous study of darter diving behaviour reported no relationship between dive duration and water depth, contrary to optimal dive models. In this study I provide more extensive observations of African darters Anhinga melanogaster rufa diving in water

SUMMARY 1. We tested the hypothesis that indirect food web interactions between some common, invertivorous fishes and their prey would positively affect growth of an algivorous fish species. Specifically, we predicted that orangethroat darter (Etheostoma spectabile) would increase periphyton biomass via a top-down pathway, indirectly enhancing growth of the algivorous central stoneroller minnow (Campostoma anomalum). Moreover, we predicted that sand shiner (Notropis stramineus) would increase periphyton biomass via a bottom-up pathway and indirectly enhance growth of the stoneroller minnow. 2. In an 83-day experiment in large, outdoor, stream mesocosms, we stocked two fish species per mesocosm (stoneroller and either darter or shiner), estimated the effects of the invertivorous and grazing fishes on periphyton biomass and estimated growth of the algivorous fish. 3. The darter consumed grazing invertebrates, indirectly increasing periphyton biomass. The shiner consumed terrestrial insects as predicted, but it did not affect periphyton biomass. 4. In support of our hypothesis, the darter indirectly enhanced stoneroller growth. As predicted, stonerollers consumed the increased periphyton in streams with darters, resulting in greater growth, condition and gut fullness compared to streams without darters. No indirect interaction was observed between stonerollers and shiners. 5. Our study suggests that some invertivorous fish species can positively affect growth of algivorous fishes through indirect food web interactions. Thus, in stream communities, it is possible that the loss of a single, invertivorous fish taxon could have negative consequences on algivorous fish populations via the removal of positive indirect food web interactions.

We used histological features to identify the spawning chronologies of river-dwelling populations of slenderhead darter Percina phoxocephala, suckermouth minnow Phenacobius mirabilis, stonecat Noturus flavus, and red shiner Cyprinella lutrensis and to relate their reproductive status to microhabitat associations. We identified spawning and nonspawning differences in habitat associations resulting from 1 year of field data via logistic regression modeling and identified shifts in microhabitat selection via frequency-of-use and availability histograms. Each species demonstrated different habitat associations between spawning and nonspawning periods. The peak spawning period for slenderhead darters was April to May in high-velocity microhabitats containing cobble. Individuals were associated with similar microhabitats during the postspawn summer and began migrating to deeper habitats in the fall. Most suckermouth minnow spawned from late March through early May in shallow microhabitats. The probabil...