Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

Patch selection by mottled sculpin (Pisces: Cottidae) in a southern Appalachian stream

1. We examined the effects of prey abundance on patch selection by a benthic fish, the mottled sculpin ( Cottus bairdi ), in a fourth order, southern Appalachian stream (North Carolina, U.S.A.). This habitat is a mosaic of small (i.e. < 0.5 m²), relatively discrete patches.
2. Patches were characterized in terms of physical habitat variables, detritus (coarse particulate organic matter, CPOM) and macroinvertebrate abundance (number and biomass). We quantified patch selection by comparing the characteristics of patches utilized by sculpin with those of locally available patches. Locally available patches were selected using a constrained random sampling design (i.e. randomly selected within a 2 m radius from each fish). We also examined the relationship between macroinvertebrate abundance, CPOM and the physical characteristics of available patches.
3. Patches selected by sculpin contained significantly higher macroinvertebrate abundances (both number and biomass) than locally available patches in five out of six seasonal samples. Sculpin also occupied patches with significantly higher amounts of CPOM in three out of five seasonal samples. Patches utilized by sculpin, however, could not be consistently differentiated from locally available patches on the basis of physical variables. In addition, macroinvertebrate abundance was not consistently related to physical habitat variables or CPOM during the course of the study.
4. Our results suggest that sculpin are able to assess patch quality on the basis of prey abundance and select patches that potentially maximize energy gain. This behaviour may produce an increase in individual fitness, especially when prey distributions are not consistently related to habitat variables. Quantifying patch use in relation to prey abundance may help elucidate the causal factors determining habitat use by benthic fishes in other lotic systems.     

Patch selection by mottled sculpin (Pisces: Cottidae) in a southern Appalachian stream

1. We examined the effects of prey abundance on patch selection by a benthic fish, the mottled sculpin ( Cottus bairdi ), in a fourth order, southern Appalachian stream (North Carolina, U.S.A.). This habitat is a mosaic of small (i.e. < 0.5 m²), relatively discrete patches.
2. Patches were characterized in terms of physical habitat variables, detritus (coarse particulate organic matter, CPOM) and macroinvertebrate abundance (number and biomass). We quantified patch selection by comparing the characteristics of patches utilized by sculpin with those of locally available patches. Locally available patches were selected using a constrained random sampling design (i.e. randomly selected within a 2 m radius from each fish). We also examined the relationship between macroinvertebrate abundance, CPOM and the physical characteristics of available patches.
3. Patches selected by sculpin contained significantly higher macroinvertebrate abundances (both number and biomass) than locally available patches in five out of six seasonal samples. Sculpin also occupied patches with significantly higher amounts of CPOM in three out of five seasonal samples. Patches utilized by sculpin, however, could not be consistently differentiated from locally available patches on the basis of physical variables. In addition, macroinvertebrate abundance was not consistently related to physical habitat variables or CPOM during the course of the study.
4. Our results suggest that sculpin are able to assess patch quality on the basis of prey abundance and select patches that potentially maximize energy gain. This behaviour may produce an increase in individual fitness, especially when prey distributions are not consistently related to habitat variables. Quantifying patch use in relation to prey abundance may help elucidate the causal factors determining habitat use by benthic fishes in other lotic systems.     

Abundance and distribution within a guild of benthic stream fishes: local processes and regional patterns

1. Spatial patterns at regional and local scales were examined for evidence that species interactions can influence distribution and abundance within a guild of benthic fishes in upland streams of Oklahoma, U.S.A. Three groups of community patterns were examined: the species–area relationship, species–habitat associations, and interspecific associations.
2. The species–area relationship for riffle habitats was compared to a null species–area model based on random placement. The observed species–area curve was steeper resulting in less species per unit area in small streams than predicted by the null model.
3. Small, species-poor streams had summed fish densities at least as high as larger, species-rich streams, suggesting density compensation.
4. Several significant patterns of negative covariation were found among species at the regional scale, before and after statistically accounting for effects due to measured habitat variables.
5. For two of these negatively covarying taxa ( Cottus carolinae and Etheostoma spectabile ), the influence of each species on the distribution of the other was evaluated experimentally in field enclosures varying in depth and current velocity. The sculpin C . carolinae caused a shift in habitat use by the darter E . spectabile , but no reciprocal shift was found.
6. These results indicate an agreement between local and regional patterns of distribution for C . carolinae and E . spectabile and suggest that biotic interactions can influence regional patterns of distribution for species within this guild.     

Which species,how many,and from where: Integrating habitat suitability,population genomics,and abundance estimates into species reintroduction planning

Extirpated organisms are reintroduced into their former ranges worldwide to combat species declines and biodiversity losses. The growing field of reintroduction biology provides guiding principles for reestablishing populations, though criticisms remain regarding limited integration of initial planning, modeling frameworks, interdisciplinary collaborations, and multispecies approaches. We used an interdisciplinary, multispecies, quantitative framework to plan reintroductions of three fish species into Abrams Creek, Great Smoky Mountains National Park, USA. We first assessed the appropriateness of habitat at reintroduction sites for banded sculpin (Cottus carolinae), greenside darter (Etheostoma blennioides), and mottled sculpin (Cottus bairdii) using species distribution modeling. Next, we evaluated the relative suitability of nine potential source stock sites using population genomics, abundance estimates, and multiple‐criteria decision analysis (MCDA) based on known correlates of reintroduction success. Species distribution modeling identified mottled sculpin as a poor candidate, but banded sculpin and greenside darter as suitable candidates for reintroduction based on species‐habitat relationships and habitats available in Abrams Creek. Genotyping by sequencing revealed acceptable levels of genetic diversity at all candidate source stock sites, identified population clusters, and allowed for estimating the number of fish that should be included in translocations. Finally, MCDA highlighted priorities among candidate source stock sites that were most likely to yield successful reintroductions based on differential weightings of habitat assessment, population genomics, and the number of fish available for translocation. Our integrative approach represents a unification of multiple recent advancements in the field of reintroduction biology and highlights the benefit of shifting away from simply choosing nearby populations for translocation to an information‐based science with strong a priori planning coupled with several suggested posteriori monitoring objectives. Our framework can be applied to optimize reintroduction successes for a multitude of organisms and advances in the science of reintroduction biology by simultaneously addressing a variety of past criticisms of the field.     

Microhabitat use in a stream fish assemblage

We examined microhabitat use among fishes in a 37-m section of Coweeta Creek, North Carolina. Numerical abundances of species changed substantially during the 17-month study period. Microhabitat availability, however, did not change markedly during our investigation. Seasonal principal component analyses of microhabitat availability and fish microhabitat use data extracted two main patterns of non-random microhabitat use. Clinostomus funduloides, Notropis eoccogenis and Semotilus atromaculatm were significantly over-represented in deep areas with low to intermediate velocities and depositional substrata. Campostoma oligolepis, Coitus bairdi, Etheostoma blennioides, Rhinichthys cataractae and Salmo gairdneri all occurred in intermediate to deep microhabitats with moderate to high velocities and erosional substrata. Five of seven species exhibited seasonal variation in microhabitat utilization, whereas six species displayed size-related variation in use. Size-related variation was probably ontogenetic. We attributed most seasonal changes in microhabitat use to variations in microhabitat availability.
We used canonical analysis of discriminants to identify factors maximizing interspecific differences in microhabitat use. This analysis indicated that species could be assigned to either a benthic or a water column guild. Species within a guild generally could not be differentiated statistically, whereas members of different guilds were readily separable. These patterns persisted throughout the study, despite changes in numerical abundances of assemblage members. There was no evidence of either exploitation or interference competition for microhabitat, consequently it is unlikely that spatial resources were limiting during our study.     

A narrow hybrid zone between two Cottus species in Wills Creek, Potomac drainage

We describe a narrow hybrid zone between the mottled sculpin (Cottus b. bairdi) and the Blue Ridge sculpin (C. caeruleomentum). Seven characters (dorsal fin rays, pectoral fin rays, caudal base band condition, male spawning coloration, and one frequency and two fixed allozyme differences) distinguish the two taxa in the hybrid zone. C. caeruleomentum and C. b. bairdi diverged in these characters in allopatry as indicated by their distribution on opposite sides of the Atlantic-Ohio divide. However, a stream capture placed these two taxa in secondary contact in Wills Creek, Potomac drainage (Atlantic slope). Allozyme data indicate the presence of post-F(1) hybrids in the zone of secondary contact. Changes in allozymes, morphology, and spawning coloration along a transect in Wills Creek reveal the hybrid zone is less than 20 river kilometers in length. Estimates of root mean square dispersal and gene flow tentatively suggest that selection is operating in the Wills Creek hybrid zone. C. b. bairdi and C. caeruleomentum are maintaining their identity in seven distinguishing characters on opposite ends of the hybrid zone revealing these two taxa are independent evolutionary lineages.     

Brown trout and food web interactions in a Minnesota stream

1. We examined indirect, community‐level interactions in a stream that contained non‐native brown trout (Salmo trutta Linnaeus), native brook trout (Salvelinus fontinalis Mitchill) and native slimy sculpin (Cottus cognatus Richardson). Our objectives were to examine benthic invertebrate composition and prey selection of fishes (measured by total invertebrate dry mass, dry mass of individual invertebrate taxa and relative proportion of invertebrate taxa in the benthos and diet) among treatments (no fish, juvenile brook trout alone, juvenile brown trout alone, sculpin with brook trout and sculpin with brown trout). 2. We assigned treatments to 1 m² enclosures/exclosures placed in riffles in Valley Creek, Minnesota, and conducted six experimental trials. We used three designs of fish densities (addition of trout to a constant number of sculpin with unequal numbers of trout and sculpin; addition of trout to a constant number of sculpin with equal numbers of trout and sculpin; and replacement of half the sculpin with an equal number of trout) to investigate the relative strength of interspecific versus intraspecific interactions. 3. Presence of fish (all three species, alone or in combined‐species treatments) was not associated with changes in total dry mass of benthic invertebrates or shifts in relative abundance of benthic invertebrate taxa, regardless of fish density design. 4. Brook trout and sculpin diets did not change when each species was alone compared with treatments of both species together. Likewise, we did not find evidence for shifts in brown trout or sculpin diets when each species was alone or together. 5. We suggest that native brook trout and non‐native brown trout fill similar niches in Valley Creek. We did not find evidence that either species had an effect on stream communities, potentially due to high invertebrate productivity in Valley Creek.     

Spatial variation in density of stream benthic fishes in northern Hokkaido, Japan: does riparian vegetation affect fish density via food availability?

The densities of two benthic fishes, the Siberian stone loach (Noemacheilus barbatulus) and the wrinklehead sculpin (Cottus nozawae), and the biomass of their food resources (i.e., periphyton and benthic invertebrates) were compared between forest and grassland streams in northern Hokkaido, Japan, to examine whether riparian deforestation had positive effects on the benthic fishes via enhancement of food availability. The comparisons indicated that riparian vegetation had little influence on periphyton, invertebrates, or fishes. Regression analysis indicated that spatial variations in loach and sculpin densities were explained more by substrate heterogeneity, competitor abundance, or both, rather than by food abundance. However, when the two species were combined as benthic insectivores, a strong correlation was found between total benthic fish density and invertebrate biomass. Our results suggest that, although total benthic fish abundance was food limited, riparian vegetation had no positive effects via food availability on the benthic fishes in our streams.     

Behavioral effects of invaders: alien crayfish and native sculpin in a California stream

Invasive crayfish have been shown to have negative impacts on a range of taxa, though the mechanisms for those effects have not always been evaluated. Signal crayfish (Pacifastacus leniusculus) in Sagehen Creek were associated with reduced growth rates and gut fullness of Paiute sculpin (Cottus beldingi) in earlier experiments. This paper assesses potential behavioral mechanisms of competition between the two species. I conducted experiments to determine crayfish effects on sculpin behavior and habitat use in a stream observation facility at the Sagehen Research Station, California, USA. Sculpin reduced their use of refuges and pools, shifted into higher-velocity microhabitats, and spent more time fleeing in the presence of crayfish. Crayfish used refuges, pools, and low-velocity habitats more than sculpin in either treatment. Both species were notably nocturnal, with most activity at dusk and night observations, although crayfish were more strongly so than sculpin. Detailed field surveys of lower Sagehen Creek found that potential refuges (unembedded rocks) were closely associated with total crayfish and sculpin numbers, suggesting that cover is at least sometimes limiting under natural conditions. By displacing sculpin from refuges and pools and increasing their activity rate, crayfish may increase the likelihood of predation on sculpin. Behavioral shifts in sculpin appear to be at least partly responsible for the reduced growth rates of sculpin in the presence of crayfish.     

Comments on the morphology of Nicsmirnovius eximius (Kiser, 1948) (Aloninae,Anomopoda, Cladocera) in Thailand,with description of its male

The density, diet and habitat use of brown trout (Salmo trutta) and Siberian sculpin (Cottus poecilopus) were studied in the subalpine River Atna in southeastern Norway in the autumn during a six year period (1986–1991). There was an inverse relationship between the density of brown trout and Siberian sculpin. Diet overlap, as indicated by the Schoener index, was high between the two species, ranging between 0.48 and 0.86. Chironomid larvae and other aquatic insects were the most common food items for both species. Brown trout also consumed substantive amounts of surface insects. Siberian sculpin typically occupied sites with finer substrates and greater water depths than brown trout, even though there was considerable overlap in habitat use between the two species. Because the two species shared similar habitats, we suggest that the potential for species interactions exists, particularly at sites where density of sculpin is high.     

Habitat partitioning in a community of darters in the Elk River, West Virginia

Microhabitat use and habitat partitioning for 10 darter species (Percidae: Etheostomatini) were studied by direct observation (snorkeling) at five sites in the Elk River drainage, West Virginia, U.S.A. We used canonical discriminant analysis and univariate tests to determine microhabitat variables that were important in segregating species. Darter species were segregated by depth, substrate size, and water velocity. In addition, Percina typically occurred in the water column, whereas species of Etheostoma were benthic and segregated by occurring under, between, and on top of rocks.     

Learning to Avoid Dangerous Habitat Types by Aquatic Salamanders,Eurycea tynerensis

There should be intense selection for predation avoidance mechanisms when prey live in close proximity to their predators. Prey individuals that can learn to associate habitat features with high levels of predation risk should experience increased survival if they subsequently avoid those habitats. We tested whether or not habitat learning occurred in a benthic stream community consisting of adult Oklahoma salamander (Eurycea tynerensis) prey and a syntopic predatory fish, the banded sculpin (Cottus carolinae). We exposed individual salamanders to chemical stimuli from sculpin, non‐predatory tadpoles, or a blank control in training tanks containing either rocks or grass. Two days later, the salamanders were tested in tanks that offered a choice of rocks or grass. Salamanders showed significant avoidance of the habitat where they had previously encountered chemical cues from sculpin in comparison to the non‐predatory controls. Learning to avoid dangerous habitats may be particularly important for prey whose predators are visually cryptic ambush foragers, such as sculpin.     

Comparative ecology of prickly sculpin,Cottus asper,and coastrange sculpin,C. aleuticus,in the Eel River,California

Synopsis We documented species' distributions, size structure of populations, abundance in mainstem and tributary streams, habitat use, and diets of prickly sculpin, Cottus asper, and coastrange sculpin, C. aleuticus, in the Eel River drainage of California, to determine the processes allowing coexistence of these very similar fishes. We observed prickly sculpins at 43 sites and coastrange sculpins at 34. The species co-occurred at 26 sites. Young-of-year coastrange sculpins were only observed within 42 km of the ocean, but young-of-year prickly sculpins were present throughout the species range. Mean, maximum, and minimum lengths of coastrange sculpins were positively correlated with distance from the ocean but no significant relationships were found for prickly sculpins. Absolute abundance of both species was highest in mainstem habitat (prickly sculpins = 0.6 sculpins m^–2 and coastrange sculpins = 0.4 sculpins m^–2) . Tributary densities of both species tended to be less than 0.1 sculpins m^–2. The species inhabited very similar habitats and had very similar diets. Coastrange sculpin populations in upstream areas were maintained by immigration from downstream areas in contrast with prickly sculpin populations that produced young-of-year fish throughout their range. Densities were probably not high enough for interspecific interactions to be important. The factors limiting the upstream distribution of the species may include high water temperatures, stability of the stream bed, and behavior of the fish. In the past, the range of sculpins within the Eel River drainage probably fluctuated with changing physical conditions. Recent introductions of exotic species that compete with and prey upon sculpins, and ongoing human activities in the drainage could result in major reductions in the distribution and abundance of one or both species.     

Zebra mussels affect benthic predator foraging success and habitat choice on soft sediments

The introduction of zebra mussels (Dreissena spp.) to North America has resulted in dramatic changes to the complexity of benthic habitats. Changes in habitat complexity may have profound effects on predator-prey interactions in aquatic communities. Increased habitat complexity may affect prey and predator dynamics by reducing encounter rates and foraging success. Zebra mussels form thick contiguous colonies on both hard and soft substrates. While the colonization of substrata by zebra mussels has generally resulted in an increase in both the abundance and diversity of benthic invertebrate communities, it is not well known how these changes affect the foraging efficiencies of predators that prey on benthic invertebrates. We examined the effect of zebra mussels on the foraging success of four benthic predators with diverse prey-detection modalities that commonly forage in soft substrates: slimy sculpin (Cottus cognatus), brown bullhead (Ameirus nebulosus), log perch (Percina caprodes), and crayfish (Orconectes propinquus). We conducted laboratory experiments to assess the impact of zebra mussels on the foraging success of predators using a variety of prey species. We also examined habitat use by each predator over different time periods. Zebra mussel colonization of soft sediments significantly reduced the foraging efficiencies of all predators. However, the effect was dependent upon prey type. All four predators spent more time in zebra mussel habitat than in either gravel or bare sand. The overall effect of zebra mussels on benthic-feeding fishes is likely to involve a trade-off between the advantages of increased density of some prey types balanced against the reduction in foraging success resulting from potential refugia offered in the complex habitat created by zebra mussels.     

Microhabitat partitioning in a diverse assemblage of darters in the Allegheny River system

Synopsis Habitat partitioning among eleven species of darters (Percidae: Etheostomatini) from the Allegheny River system was studied through underwater observation. Percina caprodes and Percina copelandi showed consistent segregation from Etheostoma by occupying deeper habitats. Substrate size, depth, and water velocity were important variables by which Etheostoma species segregated. Analysis of niche breadth values indicated that species differed widely in their degree of specialization in habitat use; based on the variables measured, Etheostoma zonale was a habitat generalist whereas Etheostoma camurum, Etheostoma tippecanoe, and Percina caprodes tended towards habitat specialization. Habitat segregation appears to be an important mechanism allowing the coexistence of these closely related and ecologically similar species. Microhabitat quantification on a fine scale was important in discovering habitat differences in this diverse system.     

Habitat use and its implications to functional morphology: niche partitioning and the evolution of locomotory morphology in Lake Tanganyikan cichlids (Perciformes: Cichlidae)

Animal locomotory morphology, i.e. morphological features involved in locomotion, is under the influence of a diverse set of ecological and behavioral factors. In teleost fish, habitat choice and foraging strategy are major determinants of locomotory morphology. In this study, we assess the influence of habitat use and foraging strategy on important locomotory traits, namely the size of the pectoral and caudal fins and the weight of the pectoral fin muscles, as applied to one of the most astonishing cases of adaptive radiation: the species flock of cichlid fishes in East African Lake Tanganyika. We also examine the course of niche partitioning along two main habitat axes, the benthic vs. limnetic and the sandy vs. rocky substrate axis. The results are then compared with available data on the cichlid adaptive radiation of neighbouring Lake Malawi. We find that pectoral fin size and muscle weight correlate with habitat use within the water column, as well as with substrate composition and foraging strategies. Niche partitioning along the benthic–limnetic axis in Lake Tanganyikan cichlids seems to follow a similar course as in Lake Malawi, while the course of habitat use with respect to substrate composition appears to differ between the cichlid assemblages of these two lakes.     

Habitat use and feeding behavior of thirteen species of benthic stream fishes

Synopsis The densities, habitat use, and feeding behaviors of 13 fish species belonging to the benthic invertebrate-feeding guild were studied by snorkeling at three localities in the Little River of eastern Tennessee, U.S.A. Resource partitioning occurred by habitat, feeding behavior and time of activity. Differences were also found at the generic level.Cottus was a nocturnal feeder, whereasPercina andEtheostoma were, for the most part, diurnally active.Percina moved about rapidly and spent most of its time above the bottom. In contrast,Etheostoma varied considerably in the amount of time spent under cover, spent little time above the bottom, and exhibited low levels of swimming activity. Nearly all species sought cover at night, suggesting they may be particularly sensitive to predation at night. Species with small adult sizes (Etheostoma, Cottus andP. evides) were concentrated in shallow water habitats, whereas species with large adult sizes (Percina) were more abundant in deep water habitats. The habitat use data are consistent with the hypothesis that size-selective predation by centrarchid bass may cause smaller fish to avoid deep water areas. Large species should have a lower risk of predation due to their size and behavior.     

Patch based predation in a southern Appalachian stream

Streams are characterized by high degrees of patchiness that could influence the role of predators in these systems. Here we assess the impact of predatory benthic fishes on benthic macroinvertebrate density, biomass, and community structure at the patch scale in a fourth order stream in the southern Appalachians. We tested the role of predation in two different patch types: patches inhabited by adult mottled sculpin ( Cottus bairdi ) and random patches. We placed 30 basket pairs (one open to fish predation, and one from which fish predators were excluded) in the streambed at each patch type. We also tested for potential basket effects by setting up a basket control area. Although there was some evidence of basket artifacts on macroinvertebrate density in sculpin patches, these artifacts were not consistent and we do not feel that they affected our results because predators did not affect macroinvertebrate density. In random patches, predation did not significantly affect macroinvertebrate density or biomass. Predators significantly reduced macroinvertebrate biomass in sculpin patches but did not affect prey density. When the data-set was size-limited to exclude macroinvertebrates too large for consumption by sculpin, macroinvertebrate biomass did not differ significantly between exclusion and open baskets. This suggests that sculpin can reduce macroinvertebrate biomass through a combination of consumption and by predator-induced emigration of large macroinvertebrates into areas that are protected from sculpin. In addition, invertebrate predator biomass was higher in predator exclusion baskets in sculpin patches indicating that predation pressure remained high in the exclusion baskets despite fish exclusion. These results illustrate the heterogeneity of streams and the effect of small-scale differences (e.g. location of predators' territories) on local processes. Experiments that utilize these differences can provide insights into these stream processes.     

Morphology and habitat use by fishes of the Rio das Velhas basin in southeastern Brazil

We analyzed the morphology and habitat use of 16 small-sized fish species that are abundant in the Rio das Velhas basin using 17 morphological attributes. Habitat use was characterized in terms of the species mean density considering three hydraulic factors: substrate, water depth and mean water velocity. The distribution of species within the morphological space demonstrated congruence between the morphological attributes of a species, and their occupation of vertical microhabitats. Based on this data, the species could be classified as benthic A, benthic B or nektonic zone-dwelling species. Benthic A and B species differed mainly in relation to their body shapes, which were fusiform or depressed, respectively. Generally, habitat selection by each species was related to certain hydraulic factors, water velocity in particular. Siluriformes generally had morphological or behavioral adaptations to live in fast flowing habitats, and they were found more frequently than Characiformes in microhabitats with greater water velocities. Benthic species were the most selective with respect to substrate type. Between congener species, the habitat use of Serrapinnus heterodon and Serrapinnus piaba greatly overlapped, but these species were separated in morphological space. In contrast, Characidium species were closely related in morphological space but had great habitat segregation. Fish requirements and habitat use are poorly understood but are important to species management and habitat restoration.