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.     

Early development of johnny darter,Etheostoma nigrum,and fantail darter,E. flabellare,with a discussion of its ecological and evolutionary aspects

Synopsis Fantail darters,Etheostoma flabellare, have larger eggs with greater yolk volume than johnny darters,E. nigrum, and have a longer embryo period. This longer embryo period is associated with accelerated finfold differentiation, producing a larger, better formed individual at the onset of exogenous feeding. Developmental differences among these two species and the rainbow darter,Etheostoma caeruleum, primarily in timing of ontogenetic events, are described in detail. These differences in timing are considered largely early developmental phenomena as they affect embryos, larvae, and aievins but apparently not juveniles or adults. Demographic aspects of early life history were assessed using data from studies of Ontario streams. Differences in early survival among species appear to play an important role in recruitment of mature females, but cannot be attributed to parental care or egg size differences.     

Early development of the northern logperch,Percina caprodes semifasciata,according to the theory of saltatory ontogeny

Synopsis The early development of northern logperch, Percina caprodes semifasciata, was examined from an ecological perspective and in relation to the theory of saltatory ontogeny. Steps, the intervals of relative homeostasis, separated by thresholds, rapid switches to new form and function resulting from canalization of changes in the preceeding step, are described and related to environmental factors. Adaptive shifts in growth allometries provided further evidence of the saltatory nature of ontogeny. Logperch eggs are small (1.2 mm diameter), demersal, and adhesive. The simple embryonic respiratory system reflects high oxygen availability on the lotic spawning grounds. Hatching occurs early, relative to other darters, and free embryos are pelagic. The small free embryos presumably drift downstream to lentic areas, where small planktonic food is more abundant. Logperch are therefore nonguarding, open substrate spawning lithopelagophils and, along with other Percina species, have not advanced from ancestral percid reproductive styles. Percina species have generally remained in ancestral habitats-rivers and lakes. The close correlation between developmental patterns of logperch and environmental factors is therefore interpreted as a restriction based on retention of ancestral characteristics, not as evidence of novel adaptations to new habitats.     

Alarm pheromone recognition in a Missouri darter assemblage

The a priori null hypotheses that responses to donor, orangethroat darter Etheostoma spectabile , johnny darter Etheostoma nigrum , logperch Percina caprodes , slenderhead darter Percina phoxocephala and blackside darter Percina maculata , extracts would be absent in the three test species, orangethroat darter, johnny darter and logperch, was rejected. This was because orangethroat and johnny darter responded with decreased activity to skin extracts derived from conspecifics and from other members of the assemblage. In contrast, logperch were notable by their lack of a detectable response to any species. Similarities and differences in habitat utilization are seen as a possible basis for these responses.     

Microsatellite markers for the endangered Roanoke logperch, Percina rex (Percidae) and their potential utility for other darter species

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.     

Rainbow Darter (Etheostoma caeruleum,Storer, 1845) predation on early ontogenetic stages of Lake Sturgeon (Acipenser fulvescens,Rafinesque, 1817)

Previous molecular diet analysis identified lake sturgeon (Acipenser fulvescens, Rafinesque, 1817) DNA in the gastrointestinal tracts of stream-resident rainbow darters (Etheostoma caeruleum, Storer, 1845) during the egg incubation, free embryo, and larval drift stages. The objectives of this experimental study were to: (a) quantify levels of predation by rainbow darters on lake sturgeon at the egg and free-embryo stages; and (b) evaluate whether predation varied as a function of substrate size and rainbow darter body size. We conducted experimental trials in 23-L polycarbonate tanks 0.41 m (L) × 0.33 m (W) × 0.30 m (D) with a standardized benthic area of 0.14 m². The tanks were randomly assigned one of two different substrate size classes: large rock (51.35 mm ± 0.91 mm) or small rock (27.68 mm ± 0.57 mm). We stocked individual rainbow darter, which were deprived of feed for 48 hr, with lake sturgeon (133 individuals/m²) in each of 12 replicates per ontogenetic stage and substrate type. The number of surviving lake sturgeon was quantified following a 24-hr predation exposure period. We used a generalized linear model with a binomial distribution to assess the influence of ontogenetic stage, substrate size, and rainbow darter body size on proportional lake sturgeon survival. Predation on lake sturgeon occurred at both egg (6.25 ± 1.16 individuals, mean ± 2SE) and free embryo (3.08 ± 1.08 individuals, mean ± 2SE) stages. Egg proportional survival was generally lower than at the free embryo stage in both substrate sizes; however, free embryo proportional survival was greater in small substrate trials. Rainbow darter total length did not affect the probability of lake sturgeon survival at either developmental stage. Results demonstrate that rainbow darters prey on early ontogenetic stages of lake sturgeon, corroborating previous results based on genetic diet analysis. Results fill a major knowledge gap concerning the vulnerability of pre-drift sturgeon to predation by an abundant river resident species that was previously discounted as a predator for early ontogenetic stages of lake sturgeon due to its small body size.     

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.     

Seasonal changes in low oxygen tolerance of fantail,Etheostoma flabellare,rainbow, E. caeruleum,and greenside,E. blennioides,darters

Synopsis The tolerance of fantail, rainbow, and greenside darters to low oxygen was determined by exposing fish to conditions of progressively decreasing oxygen levels and recording the oxygen concentration at which loss of equilibrium occurred. The greenside darter was least tolerant to low oxygen levels in summer, with loss of equilibrium occurring at a mean O₂ concentration of 3.39 mg l⁻¹. Tolerances then increased through spring. A similar trend was observed for the fantail darter, with the lowest tolerance occurring in summer (2.36 mg ⁻¹). and greatest tolerance in spring (2.03 mg l⁻¹). The rainbow darter exhibited its greatest tolerance to low oxygen in summer and spring (1.64 and 1.93 mg l⁻¹, respectively). Interspecific differences were greatest in summer, with greenside darters being the least tolerant to low oxygen levels and the rainbow darter being the most tolerant. No differences in tolerance among the species were noted in winter. The low oxygen tolerance exhibited in summer by the greenside darter may be due to a higher metabolic cost and a higher critical oxygen tension at maximal stream temperatures (>25° C). The observed differences in oxygen tolerance may influence the longitudinal distribution and habitat selection of darters, especially in the summer months.     

Patterns of shield darter, <Emphasis Type="Italic">Percina peltata</Emphasis>, distribution in the Eastern Piedmont of Maryland,USA

Detailed analyses of habitat associations with rare species are typically constrained by limited sample size and the availability of habitat data. The dense spatial coverage of stream sampling by the Maryland Biological Stream Survey provides ample data to quantitatively examine correlations between habitat and rare species distributions. The shield darter, Percina peltata, has a widespread distribution on the Atlantic Slope of the United States, but is uncommon throughout its range in Maryland. Associations of in situ physical habitat, water chemistry, and alterations in landscape with shield darter presence in the Eastern Piedmont physiographic province in Maryland were examined. Shield darter occurrence was associated with larger sized streams in concordance with the species’ known ecology. Shield darter distribution was further associated with stream segments with deep riffle habitats with diverse velocities, low concentrations of chloride and sulfate, low levels of urbanization in upstream catchments, and several pollution intolerant fish species. Although the exact mechanism of the effects is not clear, results indicate that the shield darter is sensitive to urban development and habitat and water quality alteration that typically accompanies urbanization. Shield darter conservation in Maryland necessitates the protection and restoration of minimally urbanized watersheds where they are known to occur. The results from this study indicate that habitat information on rare species may be important in elucidating important habitat associations that are not evident via examination of community level data.     

Nest defense and aggressive interactions between a small benthic fish (the johnny darter Etheostoma nigrum) and crayfish

Synopsis Aggression by nest-guarding male johnny darters, Etheostoma nigrum, against intruding crayfish was investigated in laboratoy experiments and field observations. In the laboratory, darter success in chasing crayfish, Orconectes rusticus, from the nest site was inversely related to crayfish size. Small crayfish (less than 15 mm carapace length) were routinely evicted from the nest area by nips directed at the posterior end of the abdomen. Although such aggressive behavior was less successful against larger crayfish, even the largest crayfish tested (carapace length 30–32 mm) were chased from the nest area in 33% of the trials. Those large crayfish that entered nests often remained despite repeated attacks by the male johnny darter and egg predation was observed. In a small Ohio stream, openings to johnny darter nests were generally between 7 and 13 mm. Thus crayfish with a carapace height greater than 13 mm (corresponding to a carapace length greater than 29 mm) would not be able to enter johnny darter nests. In field observations, male Johnny darters successfully defended nests against another crayfish species (Orconectes sanborni, carapace length 12–29 mm). Together, aggressive behavior and small size of nest entrances allow the johnny darter to successfully reproduce in areas with abundant crayfish. The Unit is sponsored jointly by the United States Fish and Wildlife Service, The Ohio Department of NaturalResources, and The Ohio State University.     

The Round Goby, Neogobius melanostomus, a Fish Invader on both sides of the Atlantic Ocean

During the past decade, a bottom-dwelling, aggressive, multiple-spawning fish, the round goby (Gobiidae: Neogobius melanostomus), has spread from its native region in the Ponto-Caspian throughout Europe and to the Laurentian Great Lakes in North America. An international workshop, held at the Hel Marine Station, Poland, was organized to summarize population features of the round goby. Common fish predators of round gobies in the Great Lakes and in native regions are obligate and facultative benthic fishes and occasionally, pelagic fishes. In contrast, the main predator of the round goby in the Gulf of Gdansk is the Great Cormorant (Phalacrocorax carbo). In the Great Lakes, round gobies have lead to the decline of mottled sculpin (Cottus bairdi) and logperch (Percina caprodes) and reduced the hatching success of native fishes by feeding on their eggs. In the Gulf of Gdansk, round gobies have increased in abundance, while three-spined sticklebacks (Gasterosteus aculeatus) have declined. Round gobies have a broad diet throughout their range; larger specimens are molluscivores. There are fewer species of parasites and lower infection rates of round gobies in recently colonized areas than in native areas. Overall, newly colonized round gobies in brackish waters and lakes are smaller, mature earlier, have a male biased operational sex ratio and are more short-lived compared with round gobies from marine (native) habitats.     

Egg viability decreases rapidly with time since ovulation in the rainbow darter Etheostoma caeruleum: implications for the costs of choosiness

Egg viability in the rainbow darter Etheostoma caeruleum, a fish apparently lacking female mate choice, was found to decline rapidly after ovulation. It was observed that the majority of a female's clutch may fail to hatch if she is prevented from mating for as little as 6 h. These data suggest that exercising female mate preferences may be selectively disfavoured in E. caeruleum due to the high cost of delaying mating.     

Reproductive isolation between two darter species is enhanced and asymmetric in sympatry

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.     

Expression of the oligopeptide transporter, PepT1, in larval Atlantic cod (Gadus morhua)

The intestinal absorption of di- and tri-peptides generally occurs via the oligopeptide transporter, PepT1. This study evaluates the expression of PepT1 in larval Atlantic cod (Gadus morhua) during the three weeks following the onset of exogenous feeding. Larval Atlantic cod were fed either wild captured zooplankton or enriched rotifers. cDNA was prepared from whole cod larvae preceding first feeding and at 1000 each Tuesday and Thursday for the following three weeks. Spatial and temporal expression patterns of PepT1 mRNA were compared between fish consuming the two prey types using in situ hybridization and quantitative real-time PCR. Results indicated that PepT1 mRNA was expressed prior to the onset of exogenous feeding. In addition, PepT1 was expressed throughout the digestive system except the esophagus and sphincter regions. Expression slightly increased following first-feeding and continued to increase throughout the study for larvae feeding on both prey types. When comparing PepT1 expression in larvae larger than 0.15-mg dry mass with expression levels in larvae prior to feeding, no differences were detected for larvae fed rotifers, but the larvae fed zooplankton had significantly greater PepT1 expression at the larger size. In addition, PepT1 expression in the zooplankton fed larvae larger than 0.15-mg dry mass had significantly greater expression than rotifer fed larvae of a similar weight. Switching prey types did not affect PepT1 expression. These results indicate that Atlantic cod PepT1 expression was slightly different relative to dietary treatment during the three weeks following first-feeding. In addition, PepT1 may play an important role in the larval nutrition since it is widely expressed in the digestive tract.     

Differences in morphological growth trajectories between greenside darter Etheostoma blennioides populations from two distinct drainages

The greenside darter Etheostoma blennioides is a small-bodied benthic stream fish that occurs in multiple drainages of the eastern USA. Previous studies have revealed some morphological differentiations among greensides from isolated drainage systems but growth relationships among functional morphological characteristics have not been examined within and between populations. Specifically, we tested for differences in functional morphology and allometric distinctions in morphological growth trajectories between greenside populations from two drainages, the Osage River and White River. Morphological differences between individuals of the two drainages included more dorso-ventrally compressed bodies, longer snouts and larger jaw features in White River darters. Furthermore, body depth, snout length and jaw width grow at significantly different trajectories in individuals of the two drainages. Individuals of the White River drainage (generally a higher gradient stream system) become comparatively more streamlined and develop proportionately larger feeding related traits during growth. These results suggest that individuals of both drainages share a similar morphology during early life but deviate during continued development and this may be partially due to environmentally induced plastic response. This study represents a novel approach for comparing morphological development among fish populations and presents environmental factors (e.g., stream gradient and elevation) that potentially influence functional characteristics.     

Association between parasite load and orange,but not blue,male nuptial colouration in Etheostoma caeruleum

The hypothesis that males possessing more vivid nuptial colouration are of better quality with regard to individual measures of health such as parasite infection was investigated by taking spectral measurements of orange and blue bars in the nuptial males of a sexually dichromatic stream fish rainbow darter Etheostoma caeruleum to determine if male colouration in this species varied with parasite load in the form of black spot disease. The yellow chroma of orange bars, i.e. the relative contribution of wavelengths in the range of 550–625 nm to total brightness, was the only spectral measurement significantly associated with parasite counts. These results are discussed in the context of sexual selection and the potential of orange bars in E. caeruleum to serve as honest indicators of quality to potential mates.     

Comparison of low oxygen avoidance in a bimodal breather,Erpetoichthys calabaricus and an obligate water breather,Percina caprodes

Synopsis Behavioral responses of the African reedfish, Erpetoichthys calabaricus, (Polypteridae), a bimodal breather, and the logperch, Percina caprodes, (Percidae), an obligate water breather, to low concentrations of dissolved oxygen were examined in a two-choice experimental design. In contrast to logperch, which consistently avoided dissolved oxygen concentrations of about 2.0 mg 1⁻¹, reedfish did not exhibit strong avoidance to dissolve oxygen concentrations of 0.5 mg 1⁻¹ and less.     

Biochemical, histological and behavioural aspects of visual function during early development of rainbow trout

Retinal structure and concentration of retinoids involved in phototransduction changed during early development of rainbow trout Oncorhynchus mykiss , correlating with improvements in visual function. A test chamber was used to evaluate the presence of optokinetic or optomotor responses and to assess the functionality of the integrated cellular, physiological and biochemical components of the visual system. The results indicated that in rainbow trout optomotor responses start at 10 days post‐hatch, and demonstrated for the first time that increases in acuity, sensitivity to low light as well as in motion detection abilities occur from this stage until exogenous feeding starts. The structure of retinal cells such as cone ellipsoids increased in length as photopic visual acuity improved, and rod densities increased concurrently with improvements in scotopic thresholds (2·2 log₁₀ units). An increase in the concentrations of the chromophore all‐trans‐retinal correlated with improvements of all behavioural measures of visual function during the same developmental phase.     

Diet composition of larval and young-of-year shovelnose sturgeon in the Upper Missouri River

Obtaining food following the transition from endogenous to exogenous feeding and during the first year of life is a critical event that strongly influences growth and survival of young‐of‐year fishes. For shovelnose sturgeon Scaphirhynchus platorynchus, limited information is available on food habits during the first year of life. The objective of this study was to quantify diet components of shovelnose sturgeon during the transition from endogenous to exogenous feeding and during the young‐of‐year life stage in the North Dakota and Montana portions of the Missouri River. Young‐of‐year shovelnose sturgeon were sampled between early August and early September 2003. Shovelnose sturgeon initiated exogenous feeding by 16 mm, and individuals 16–140 mm fed exclusively on two macroinvertebrate orders (Diptera and Ephemeroptera). Young‐of‐year shovelnose sturgeon exhibited an apparently high feeding success as 99 of 100 individuals contained food in the gut. The number of organisms in the gut increased exponentially with fish length for larval Diptera (r² = 0.73, P < 0.0001) and linearly (r² = 0.12, P = 0.0006) for larval Ephemeroptera, but the number of Diptera pupae in the gut was not significantly related (P = 0.55) to length of young‐of‐year shovelnose sturgeon. The length of ingested prey was linearly related to fish length for Diptera larvae (r² = 0.20, P = 0.002), whereas the relationship between lengths of ingested Ephemeroptera larvae and lengths of young‐of‐year shovelnose sturgeon was best described by a power function (r² = 0.50, P < 0.0001). These results provide the first quantification of feeding dynamics for young‐of‐year shovelnose sturgeon in a natural river environment.     

Ontogenetic shifts in habitat use by the endangered Roanoke logperch (Percina rex)

1. Conservation of the federally endangered Roanoke logperch (Percina rex, Jordan and Evermann) necessitates protection of habitat that is critical for all age classes. We examined habitat use patterns of individual logperch to determine: (1) if age classes of logperch in the Nottoway and Roanoke Rivers exhibit habitat selectivity, (2) if age classes differ in habitat use, and (3) if ontogenetic patterns of habitat use differ between the Roanoke and Nottoway river populations. 2. In the summers of 2000 and 2001, we observed 17 young‐of‐year (YOY) logperch [<4 cm total length (TL)], 13 subadult logperch (4–8 cm TL), and 49 adult logperch (>8 cm TL) in the upper Roanoke River, and 40 subadult and 39 adult logperch in the Nottoway River, Virginia. 3. All size classes of Roanoke logperch demonstrated habitat selectivity and logperch used a wide range of habitats in the Roanoke and Nottoway rivers during ontogeny. Habitat use by logperch varied among age classes and between rivers. 4. In the Roanoke River, adult and subadult logperch primarily preferred run and riffle habitat, often over gravel substrate. Subadults were found in lower water velocities and slightly more embedded microhabitats than adults. YOY logperch were found in shallow, stagnant backwaters and secondary channels. In the Nottoway River, both adult and subadult logperch were found over sand and gravel in deep, low‐velocity pools and runs. Subadults were observed in slightly more silted, lower velocity habitat than adults. Shifts in habitat use were more distinct between age classes in the Roanoke River than the Nottoway River. 5. Successful conservation of this species will involve sound understanding of spatial variation in habitat use over logperch life history and preservation of the ecological processes that preserve required habitat mosaics.