Comparative Microhabitat Use of Ecologically Similar Benthic Fishes

Although benthic insectivorous fishes such as darters and sculpins represent a significant component of riffle communities, few studies have compared the habitat use of these non-related but ecologically similar fishes. The objectives of this study were to examine the habitat use of Etheostoma olmstedi (tessellated darter) compared to Cottus bairdi (mottled sculpin) in Nescopeck Creek and Cottus cognatus (slimy sculpin) in Jack's Creek, Pennsylvania through underwater observation. Etheostoma olmstedi occupied habitats with significantly deeper waters than those available, whereas adult and young of the year Cottus occupied habitats with significantly faster water velocities than those available. Canonical discriminant analysis revealed microhabitat partitioning between E. olmstedi and each Cottus species. Cottus bairdi and C. cognatus occupied significantly shallower habitats with faster water velocities than E. olmstedi. Sculpin species were observed most frequently under substrate whereas E. olmstedi occurred most frequently on the top surface of the substrate. Hurlbert's standardized niche breadth values indicated that C. bairdi and C. cognatus were habitat specialists with regard to water velocity measures, but exhibited generalistic patterns of depth and substrate size use. Etheostoma olmstedi was a habitat specialist with respect to depth, but exhibited generalistic patterns of resource use for substrate size. Differential habitat use by these benthic fishes is consistent with the hypothesis that resource partitioning facilitates species coexistence among stream fishes.     

Microsatellite DNA markers for the rainbow darter,Etheostoma caeruleum (Percidae), and their potential utility for other darter species

The rainbow darter Etheostoma caeruleum is a small fish in the perch family (Percidae) that is adapted to fast‐flowing streams in eastern North America. It is relatively sensitive to habitat degradation and is widely used as a sentinel of stream condition. To provide a complementary tool for assessing the integrity of stream ecosystems, 16 highly polymorphic tetranucleotide microsatellite markers were identified for these darters. Between four and 16 loci were found to be useful in five congeneric species. These markers will be useful for characterizing population genetic structure and diversity of rainbow darters and related fishes.     

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.     

Spatial extent of analysis influences observed patterns of population genetic structure in a widespread darter species (Percidae)

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Comparison of visual survey and seining methods for estimating abundance of an endangered,benthic stream fish

We compared visual survey and seining methods for estimating abundance of endangered Okaloosa darters, Etheostoma okaloosae, in 12 replicate stream reaches during August 2001. For each 20-m stream reach, two divers systematically located and marked the position of darters and then a second crew of three to five people came through with a small-mesh seine and exhaustively sampled the same area. Visual surveys required little extra time to complete. Visual counts (24.2 ± 12.0; mean ± one SD) considerably exceeded seine captures (7.4 ± 4.8), and counts from the two methods were uncorrelated. Visual surveys, but not seines, detected the presence of Okaloosa darters at one site with low population densities. In 2003, we performed a depletion removal study in 10 replicate stream reaches to assess the accuracy of the visual survey method. Visual surveys detected 59% of Okaloosa darters present, and visual counts and removal estimates were positively correlated. Taken together, our comparisons indicate that visual surveys more accurately and precisely estimate abundance of Okaloosa darters than seining and more reliably detect presence at low population densities. We recommend evaluation of visual survey methods when designing programs to monitor abundance of benthic fishes in clear streams, especially for threatened and endangered species that may be sensitive to handling and habitat disturbance.     

Influence of fishes on macroinvertebrate communities and insect emergence production in intermittent stream refuges

1. Drying intermittent stream networks often have permanent water refuges that are important for recolonisation. These habitats may be hotspots for interactions between fishes and invertebrates as they become isolated, but densities and diversity of fishes in these refuges can be highly variable across time and space.
2. Insect emergence from streams provides energy and nutrient subsidies to riparian habitats. The magnitude of such subsidies may be influenced by in-stream predators such as fishes.
3. We examined whether benthic macroinvertebrate communities, emerging adult insects, and algal biomass in permanent grassland stream pools differed among sites with naturally varying densities of fishes. We also manipulated fish densities in a mesocosm experiment to address how fishes might affect colonisation during recovery from hydrologic disturbance.
4. Fish biomass had a negative impact on invertebrate abundance, but not biomass or taxa richness, in natural pools. Total fish biomass was not correlated with total insect emergence in natural pools, but orangethroat darter (Etheostoma spectabile) biomass was inversely correlated with emerging Chironomidae biomass and individual midge body size. The interaction in our models between predatory fish biomass and date suggested that fishes may also delay insect emergence from natural pools, altering the timing of aquatic–terrestrial subsidies.
5. There was an increase over time in algal biomass (chlorophyll-a) in mesocosms, but this did not differ among fish density treatments. Regardless, fish presence in mesocosms reduced the abundance of colonising insects and total invertebrate biomass. Mesocosm invertebrate communities in treatments without fishes were characterised by more Chironomidae, Culicidae, and Corduliidae.
6. Results suggest that fishes influence invertebrates in habitats that represent important refuges during hydrologic disturbance, hot spots for subsidy exports to riparian food webs, and source areas for colonists during recovery from hydrologic disturbance. Fish effects in these systems include decreasing invertebrate abundance, shifting community structure, and altering patterns of invertebrate emergence and colonisation.

     

Assessing the effects of habitat characteristics and co-occurrence with closely related species on occupancy of resident ‘wood-warblers’ in a Neotropical working landscape

The simultaneous effects of habitat traits and interspecific interactions determine the occurrence and habitat use of wildlife populations. However, little research has been devoted to examining spatial co-occurrence among closely related species while considering the effect of habitat variation and imperfect detectability of species in the field. In this study, we focused on migratory and resident 'wood-warblers' that coexist during the winter in a Neotropical working landscape in southern Mexico to understand if habitat occupancy of resident wood-warblers is influenced by habitat characteristics and by the presence of other species of resident and migratory wood-warblers. For this purpose, we implemented two-species occupancy models, which account for the imperfect detectability of these birds in the field. Our results revealed that habitat occupancy of resident wood-warblers was positively influenced by the presence of other closely related species (both migratory and resident). These positive relationships may be explained by the fact that different species of wood-warblers frequently participate in mixed-species flocks. However, these patterns of species co-occurrence were more evident among resident species than between migratory and resident species, which may be explained by micro-habitat segregation and differences in behaviours between resident and migratory wood-warblers. We also found that some habitat characteristics may mediate the observed patterns of species co-occurrence. Specifically, sites with larger trees were associated with the co-occurrence of some species of resident wood-warblers. In addition, we discuss the possibility that species co-occurrence might be the result of shared preferences for environmental factors that we did not consider. Our study highlights the importance of the interplay between species co-occurrence and habitat traits in determining the presence and habitat use of resident birds in Neotropical working landscapes.     

Positive co‐occurrence between feeding‐associative savannah fishes depends on species and habitat

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Critical current speeds and microhabitats of the benthic fishes Percina roanoka and Etheostoma flabellare

Synopsis Critical current speed (maximum current speed at which benthic stream fishes are able to hold station without active swimming) was determined in a laboratory flow chamber for Percina roanoka and Etheostoma flabellare. Mean critical current speeds differed significantly among adult P. roanoka, adult E. flabellare, and juvenile E. flabellare, and were consistent with microhabitat differences among these darters in natural environments. Behavior of E. flabellare enhanced its station-holding ability, but tests with preserved individuals suggested that morphological differences explained more of the difference in critical current speeds of the darters. On a smooth surface where grasping of the substrate was not possible, use of pectoral fins did not significantly enhance the ability of darters to maintain position in flowing water.     

Ecological and evolutionary consequences of early ontogenies of darters (Etheostomatini)

Synopsis The ecological classification of fishes into reproductive guilds is based on the premises that (1) reproductive styles and early ontogeny are closely related, and (2) both are correlated with the ecology of a species. A comparison of early ontogenies of logperch (Percina caprodes), rainbow darter (Etheostoma caeruleum), and fantail darter (E. flabellare) confirmed these premises, and provided possible explanations for diversity within the Etheostomatini. Young logperch have limited vitelline circulation, hatch while still poorly developed, and therefore must drift from oxygen rich lotic habitats to lentic habitats where small planktonic prey are available. Young rainbow and fantail darters have extensive vitelline plexuses, are well developed at transition to first feeding, and begin feeding on aquatic insects. Thus there is no necessity for a drift interval. As a result, the latter species are adapted for stream life. Interspecific differences in reproductive styles and early ontogenies may have contributed to speciation of darters by allowing partitioning of breeding sites and food resources for young. In addition, reduced drift dispersal and small stream habitation may have indirectly contributed to speciation by reducing genetic exchange among populations.     

Indirect food web interactions increase growth of an algivorous stream fish

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.     

Habitat and History Influence Abundance of Bog Turtles

Conservation of rare and endangered species requires assessment of factors that influence the current habitat associations of a species and the role of past habitat degradation in limiting occupancy or abundance. The objective of our 2011–2014 study was to determine how habitat characteristics and wetland history can predict occupancy and abundance patterns of bog turtles (Glyptemys muhlenbergii) at the fringe of their range in the southeastern United States. We used a hurdle model to examine occupancy and abundance patterns while addressing problems associated with zero-inflated data. Occupancy patterns were weakly related to percent of the wetland containing emergent vegetation, whereas abundance patterns were predicted by the percent silt in the wetland substrate, percent forest cover, amount of habitat degradation, and recovery time since past habitat degradation. The effect of historical habitat degradation on abundance rather than occupancy patterns has rarely been documented and its effect is rarely studied in vertebrate populations. Identification of predictors of occupancy and abundance patterns will aid discovery of new populations of bog turtles and improve management of occupied wetlands. © 2019 The Wildlife Society.     

Dark diversity in Amazonian stream fish communities: What factors determine species absence along environmental gradients?

1. Species distribution models often fail to predict observed patterns of species diversity, and this is because some species within a regional pool that are tolerant of conditions at a given location may nevertheless be absent from the local community. These missing species have been termed “dark diversity”. In the present study, we investigated which factors explain dark diversity among fish assemblages in Amazonian streams.
2. We sampled 71 streams in areas with different types of land use within two river basins and estimated dark diversity from patterns of species co-occurrence, using Beals’ index, along environmental gradients. From this procedure, taxa are designated as dark diversity components when they are absent from a given stream, but often co-occur with the local species at other streams, indicating similar ecological requirements. We used generalised linear models both to determine whether environmental or landscape variables, connectivity, instream environmental heterogeneity or some combination of these factors explained dark diversity of fishes, and to evaluate whether ecomorphology is associated with the extent to which a species contributes to dark diversity and which specific traits contribute the most to explaining variation in dark diversity.
3. Mean local diversity exceeded observed dark diversity. The magnitude of dark diversity was directly associated with the proportion of secondary forest in the immediate catchment and with the index of proximity to anthropogenic impact. Species that have high affinity for environments with higher current velocity, low swimming ability and that capture food mainly on the surface contributed more to dark diversity, which suggests that swimming ability, habitat preference and aspects related to diet are key predictors of the probability that a given species will be present at locations with suitable habitat.
4. Our findings reinforce the idea that dark diversity results from interactions between species traits and environmental factors, including anthropogenic impacts. Understanding the interplay among environmental factors and species traits that contribute to dark diversity provides targets for improved ecosystem restoration and sustainability of native species assemblages.

     

Impacts of co-occurring environmental changes on Alaskan stream fishes

1. Freshwater fishes are now facing unprecedented environmental changes across their northern ranges, especially due to rapid warming occurring at higher latitudes. However, empirical research that examines co-occurring environmental effects on northern fish communities remains limited.
2. We used fish community data from 1587 Alaskan stream sites to examine the potential combined and interacting effects of climate change, current weather, habitat, land use, and fire on two community-level metrics (species richness, relative abundance), and on the distributions of three Alaskan fish species.
3. Our models were 71–76% accurate in predicting the distribution of Alaskan stream fishes using a combination of climate and habitat variables. In contrast to other freshwater ecosystems that are most threatened by land use pressures, we did not detect any evidence for the potential stress of anthropogenic land use or fire on stream fishes.
4. Warming temperatures increased overall community richness and abundance but produced differing responses at the species level. Juvenile salmon presence was positively associated with several climate variables including warmer spring and autumn temperatures and wetter summers. In comparison, warmer seasonal temperatures contributed to declines for northern-adapted species such as Arctic grayling and Dolly Varden.
5. This study highlights the overarching role of current and changing climate in regulating northern stream fish biodiversity. Although many fish species may benefit from climate change across their northern ranges, localised declines are likely to occur and may prove detrimental for communities with limited fishing portfolios. Climate change adaptation and mitigation strategies customised for rapidly changing northern ecosystems will play an essential role in preserving ecologically unique northern species.

     

Reproductive strategies of Amazonian stream fishes and their fine‐scale use of habitat are ordered along a hydrological gradient

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Behavioral Isolation Based on Visual Signals in a Sympatric Pair of Darter Species

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.     

Scale-dependence of fish habitat associations in a North American river system

1. Determining the appropriate measurement scale to assess habitat variables is critical for ecologists assessing biological or ecological conditions. Depth, velocity, substrate, woody debris and other fish cover variables occur on both reach and microhabitat scales, and fish habitat associations with these variables may be scale-dependent. The aim of this work was to better understand the importance of scale for fish–habitat associations with these variables in a framework consistent with environmental filtering and to test the hypothesis that habitat variable importance is scale-dependent.
2. I used prepositioned areal electrofishing in wadeable streams of the Delaware River basin to evaluate the associations of fish with the same variables summarised on different reach and microhabitat scales. The importance of scale for fish–habitat associations was assessed using two approaches that approximate an environmental filtering framework: variance partitioning with (1) ordination and (2) generalised linear mixed models.
3. Variables on both the reach and microhabitat scales explained a significant fraction of the total variation in fish community composition (p < 0.05). Variation decomposition of reach- and microhabitat-scale effects revealed 20.2% and 2.0% of all variation were due uniquely to reach and microhabitat scales, respectively. Measures of coarseness, embeddedness, amount of riffle and areal coverage of five fish cover variables were significant explanatory variables of community composition at the reach scale only (p < 0.05). Velocity and mesohabitat (amount or presence of riffle) were the only two habitat features that were significant explanatory variables of fish community composition at both the reach and microhabitat scales (p < 0.05). Individual models of species occurrence revealed similar patterns as seen with analyses of community composition.
4. For many fishes, habitat features quantified at the reach scale were more explanatory than at the microhabitat scale. Longnose dace (Rhinichthys cataractae) were more dependent upon microhabitat variables than reach-scale variables, relative to other fishes. Mean velocity at the reach scale was the most important explanatory variable for explaining fish community composition and indicated support for the concept of environmental filtering at the reach and microhabitat scales.
5. Few studies of fish occurrence have incorporated a study design and analytical framework that approximates the hierarchical nature of habitat. This study identifies important scales and predictors, demonstrates the importance of a multiscale approach, and provides support for the environmental filtering concept at the reach and microhabitat scales. These findings will allow ecologists to better account for scale-dependent habitat associations and justify the use of fish habitat associations on reach and microhabitat scales for assessing biotic integrity, restoration and conservation of fishes.

     

Microhabitat estimation of an imperiled headwater fish,the Yazoo darter (<Emphasis Type="Italic">Etheostoma raneyi</Emphasis>), in coastal plain streams

Headwater fishes in the southeastern United States make up much of the fish biodiversity of the region yet many are imperiled. Despite this, the specific habitat requirements of imperiled headwater fishes in lowland Coastal Plain streams have rarely been quantified. Using data collected over three years of seasonal sampling we provide estimates of the microhabitat requirements of the imperiled Yazoo darter (Etheostoma raneyi Suttkus and Bart), a small benthic insectivore. Our results indicate that the species is a microhabitat specialist and that optimum microhabitat within degraded contemporary streams consists of a narrow range of water depths (about 20–30 cm), current velocity ≥ 0.25 m·s^?1, complex stable debris piles, rooted macrophytes, and likely coarse substrate. No pronounced or generalized seasonal shifts in microhabitat use occurs, and no evidence exists for intraspecific partitioning of microhabitat. Though stable and complex instream cover is one of the most important variables explaining variation in microhabitat use by Yazoo darters, such cover is rare in the degraded streams within the range of the species. Current conservation classifications of the Yazoo darter by governmental agencies and nongovernmental organizations as well as associated management plans that are based on the assumption that Yazoo darters are habitat generalists should be reviewed in recognition of the increased risk of decline because Yazoo darters are microhabitat specialists. These considerations should also be extended to other closely related imperiled species of snubnose darters.     

Spatial and temporal overlap of caracal and serval in Serengeti National Park,Tanzania

Understanding activity and habitat use are important for identifying mechanisms facilitating species co-occurrence. We studied habitat use and activity patterns of caracals (Caracal caracal) and servals (Leptailurus serval), primarily nocturnal, mid-sized felids that prey extensively on small mammals and co-occur in portions of sub-Saharan Africa. Spatial and temporal patterns of segregation were investigated in a 1,085-km² area of Serengeti National Park, Tanzania from 2010–2012. We used occupancy analysis to quantify habitat use and kernel density estimators and Mardia–Watson–Wheleer tests to analyse activity patterns. We found evidence for habitat divergence but high temporal overlap between species. Servals selected for grassland and avoided shrubland and wooded grassland. In contrast, the findings showed that caracals avoided grassland and woodland–shrubland; however, 73% of caracals were detected in wooded grassland. Overall, caracals and servals co-occurred independently, Species Interaction Factor, (phi = 1). This indicates that differential use of habitats in part facilitated coexistence of caracals and servals. Proper management of the declining grasslands including other habitats are recommended to facilitate continued coexistence. Additional studies, including feeding ecology, would be important to further understand mechanisms facilitating coexistence between caracals and servals.     

Riverscape genetics of the orangethroat darter complex

Freshwater darters belonging to the orangethroat darter species complex, or Ceasia, are widely distributed in the Central and Southern United States, with ranges that span both glaciated and unglaciated regions. Up to 15 species have been recognized in the complex, with one, Etheostoma spectabile, having a widespread northern distribution and another, Etheostoma pulchellum, having a sizeable southern distribution. The other species in the complex have much more restricted distributions in unglaciated regions of the Central Highlands. We sampled 384 darters from 52 sites covering much of the range of Ceasia and evaluated patterns of genetic diversity, genetic structure, and pre- and post-glacial patterns of range contraction and expansion. We anticipated finding much stronger signals of genetic differentiation and diversification in unglaciated regions, given the higher species diversity and levels of endemism reported there. Surprisingly, microsatellite genotyping revealed two well-differentiated genetic clusters of E. spectabile in samples from glaciated regions, one confined to the Illinois River basin and another found in the Wabash drainage and Great Lakes tributaries. This suggests that there was expansion from two isolated glacial refugia, with little subsequent post-glacial gene flow. Fish collected from throughout the unglaciated region were less genetically differentiated. Fish assigned to Etheostoma burri and Etheostoma uniporum based on collection sites and morphological characters were not genetically differentiated from E. spectabile samples from the region. Hybridization and introgression occurring in the Central Highlands may confound genetic delineation of species in this region of high endemism and diversity.