Early development of the rainbow darter,Etheostoma caeruleum,according to the theory of saltatory ontogeny

Synopsis The early development of rainbow darter, Etheostoma caeruleum, was examined from an ecological perspective. Steps and thresholds of ontogeny to completion of body squammation are defined, and related to environmental factors. Rainbow darter eggs are about 2 mm diameter, considerably larger than those of related logperch (Percina caprodes). The embryonic vitelline respiratory plexus is much more extensive. The pelagic interval characteristic of logperch and ancestral percids is eliminated and onset of exogenous feeding is delayed. The larger larvae of the rainbow darter can begin feeding directly on aquatic insects, and complete their life cycle in streams. Therefore, shifts in the timing of important thresholds (e.g. exogenous feeding) are ecologically important. Furthermore, early maturation and/or delayed bone and scale formation may be responsible for reductions in the lateralis system and scalation in this and other darter species.     

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.     

Male recognition of conspecific female chemical cues in a diverse clade of freshwater fishes

Where orangethroat darters (Etheostoma: Ceasia) and rainbow darters (Etheostoma caeruleum) co-occur, males prefer conspecific over heterospecific females. The cues males use to identify conspecific females remain unclear. We conducted behavioral trials to ask whether chemical cues function in conspecific recognition. We found that males from three orangethroat darter species preferentially associate with female scent over a control. Our results support the use of olfaction in conspecific identification in the orangethroat clade and contribute to our understanding of signals that may facilitate species recognition and underlie the evolution of behavioral isolation.     

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.     

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.     

Twenty-three microsatellite DNA loci for population genetic studies and parentage assignment in orangethroat darter, Etheostoma spectabile

The genus Etheostoma is a species-rich and ecologically important group of fishes in North America. The orangethroat darter (Etheostoma spectabile) is widely distributed and abundant in headwater streams throughout the central Midwest, and is an excellent model for ecological and mating system studies. We developed 23 novel, polymorphic, and independent microsatellite loci for E. spectabile. We found from two to 14 alleles per locus, and observed heterozygosities ranged from 0.39 to 1.0. These markers, in combination with others isolated from Etheostoma taxa, will be useful for ecological and evolutionary studies in the genus.     

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.     

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.     

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.     

Selective predation by four darter (Percidae) species on larval chironomids (Diptera) from a Mississippi stream

We determined seasonal foraging modes of four sympatric darter species, Etheostoma lynceum, E. stigmaeum, E. swaini and Percina nigrofasciata, from Beaverdam Creek, Mississippi (USA) at two scales of taxonomic resolution: (1) chironomid prey identified to family and (2) chironomid prey identified to genus/species. When chironomids were identified to family, high proportional similarity (PS) and low niche breadth (NB) values suggested the darters fed opportunistically on a relatively small number of available prey taxa. In contrast, when chironomids were identified to genus/species, concordant low PS and NB values suggested the darters fed like classic specialists, selecting a small number of prey taxa relative to prey availability in the resource base. The darters selected just one to four chironomid taxa from 52 available taxa across seasons. Our study shows that the scale of taxonomic resolution used to identify darter prey may influence the characterization of darter foraging modes.     

Distribution,abundance, and mortality of small littoral-zone fishes in Sparkling Lake,Wisconsin

Synopsis During 1982 and 1983, seining, fyke netting, and SCUBA observation were used to determine the depth distribution, abundance, and mortality of small littoral-zone fishes (bluntnose minnows, shiners [primarily mimic shiners], yellow perch, logperch, johnny darters, Iowa darters, and mottled sculpins) in Sparkling Lake, a small moderately-productive lake in northern Wisconsin. During the summer cyprinids, darters, and mottled sculpins were most abundant in areas shallower than 1 m, while yellow perch were most abundant at depths of 2 to 4 m. Between August and October cyprinids and yellow perch moved to water 1 to 2 m deeper, while the depth distributions of darters and mottled sculpins generally remained unchanged. Large within- and between-year variations in abundances and mortality rates were evident for all species. In 1982, most of the mortality of cyprinids and darters occurred during a short period in late spring and early summer, with relatively little afterwards. In 1983, this pattern was reversed for cyprinids and Iowa darters, but not logperch and johnny darters. Between-year differences in abundance were greatest for young-of-the-year yellow perch; they were 400 times more numerous in 1983 than in 1982. Darters had their lowest mortality rates and highest reproductive success in 1983, while the opposite was true for cyprinids. This lack of synchrony between darters and cyprinids suggests that these two taxa responded differently to changes in environmental conditions in Sparkling Lake. Predation may have accounted for much of the variability in darter population parameters, but appeared to be a less important source of variation for cyprinid population parameters.     

A test of three alternative pathways for consumer regulation of primary productivity

The pathways linking consumer effects to primary productivity (PPR) are likely to vary among taxa because of species-specific trophic and functional differences. Thus, it is necessary to understand the dynamics of consumer–PPR interactions so that effects of species loss on ecosystem function can be addressed from a mechanistic approach. In this study, I used three fish taxa (orangethroat darter, Etheostoma spectabile; western mosquitofish, Gambusia affinis; and bullhead minnow, Pimephales vigilax) as model consumers with different trophic and functional characteristics to test alternative mechanisms for consumer regulation of PPR (i.e., trophic cascade, terrestrial nutrient translocation, and sedimentary nutrient translocation). Experiments were conducted in stream mesocosms fitted with a combination of fish and terrestrial insect barriers to address relative importance of consumer-driven top-down and bottom-up control of PPR. A predatory invertivore, orangethroat darter, increased PPR through an apparent trophic cascade by localized reduction of benthic grazing invertebrate densities (i.e., top-down). A surface feeding insectivore, western mosquitofish, consumed terrestrial insects on the stream surface, increasing PPR by enhancing allochthonous nutrients in the mesocosms (i.e., bottom-up). A benthic omnivore, bullhead minnow, consumed benthic food items, resulting in increased PPR by enhancing availability of autochthonous nutrients via translocation of sedimentary nutrients (i.e., bottom-up). However, under specific environmental contexts, this species also consumed terrestrial invertebrates, potentially affecting PPR through terrestrial nutrient translocation as well. In this study, the trophic and functional characteristics of different species resulted in alternative pathways that increased PPR, suggesting that in natural ecosystems multiple consumer-driven pathways may be influencing PPR simultaneously and could potentially be important for temporal persistence of ecosystem function in changing environments.     

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.     

Suitability of surgically implanted 8-mm passive integrated transponder tags for small-bodied fishes

Innovative tools that benefit conservation are critical as freshwater fishes are lost at unprecedented rates. Mark-recapture methods can characterize population demographics and life-history traits of diverse fishes, but suitable techniques for tagging for individual recognition of small-bodied fishes are rare. Passive integrated transponder (PIT) tag technology may facilitate the tagging of small fishes and early life stages of larger species. However, relatively little research has evaluated the suitability of these small (8.4 × 1.4 mm) tags for many groups of small fishes. Tag loss (retention and survival) and growth of individuals implanted with PIT tags relative to control and sham treatments were compared for eight fishes of differing morphologies. Additionally, the utility of cyanoacrylate to improve tag retention was tested on a subset of taxa. Fish of each species were equally divided and randomly assigned to one of three treatment groups (handled [control], surgical incision [sham], or surgical incision and PIT tag implantation [PIT]). During the 42-d study period, mortalities and expelled tags were counted daily and growth was measured weekly. Researchers can generally expect little tag loss and uncompromised growth rates for a variety of small-bodied fishes; however, initial fish length was related to the magnitude of physiological effects for some taxa (i.e., blacknose dace Rhinichthys atratulus, johnny darter Etheostoma nigrum, juvenile white sucker Catostomus commersonii). Relatively poor survival (<80%) was observed for two benthic species: johnny darter and blacknose dace, often when incision wounds became inflamed before healing. Prevalent tag loss for johnny darter, tadpole madtom Noturus gyrinus, and a dorsal-laterally compressed cyprinid can be reduced by closing the wounds with cyanoacrylate, but with substantially increased mortality rates. This research demonstrates the broad applicability of PIT tagging technology for ecological studies of small-bodied fishes and alleviates many concerns when surgically implanting tags into several fishes.     

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.     

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.     

Territorial behaviour of the male johnny darter,Etheostoma nigrum

Synopsis A field study was undertaken to describe the territorial behaviour of male johnny darters (Etheostoma nigrum) over the reproductive cycle. While defending eggs, males performed fewer Lateral Displays and lost fewer encounters with conspecific males than in the pre-egg phase. As the reproductive cycle progressed males were less responsive to females and performed relatively fewer courtship and more aggressive displays. Nesting males apparently shifted from a sexual to a parental phase shortly after acquiring eggs.Frequency of response, responsiveness (number of responses/number of intrusions x 100) and maximum distance of response (MDR) of male johnny darters were highest early in the reproductive cycle and decreased over the egg-guarding phase. The patterns of territorial defence exhibited by male johnny darters over the reproductive cycle were similar to those of other species of fish with paternal care.     

Molecular phylogeny of the snubnose darters, subgenus Ulocentra (genus Etheostoma, family Percidae)

Snubnose darters comprise one of the largest subgenera of the percid genus Etheostoma. Many species are described based on differences in male breeding coloration. Few morphological synapomorphies have been proposed for the subgenus and their relatives, making it difficult to delineate monophyletic clades. The phylogenetic relationships of the 20 snubnose darter species of the subgenus Ulocentra and 11 members of its proposed sister subgenus Etheostoma were investigated with partial mitochondrial DNA sequences including 1033 bp encompassing the entire mitochondrial control region, the tRNA-Phe gene, and part of the 12S rRNA gene. Two hypotheses on the relationship and monophyly of the two subgenera were evaluated. Both maximum-parsimony and neighbor-joining analyses supported monophyly of the subgenus Ulocentra and resolved some species-level relationships. The banded darter, E. zonale, and its sister taxon, E. lynceum, were not closely related to the snubnose darters and appear to be diverged from the other members of the subgenus Etheostoma, fitting their former distinction as the recognized subgenus Nanostoma. The sister group to Ulocentra appears to be a restricted species assemblage within the subgenus Etheostoma containing E. blennioides, E. rupestre, E. blennius, and the E. thalassinum species group. The placement of the harlequin darter, E. histrio, is problematic, and it may represent a basal member of Ulocentra or of the restricted subgenus Etheostoma. Despite recent estimates of divergence times between nominal Ulocentra taxa, each species exhibits its own unique set of mtDNA haplotypes, providing no direct evidence for current genetic exchange between species. The nominal taxa of snubnose darters thus appear to be evolving independently from each other and therefore constitute valid species under the Phylogenetic Species Concept.     

Temperature selection and critical thermal maxima of the fantail darter,Etheostoma flabellare,and johnny darter,E. nigrum,related to habitat and season

Synopsis Riffle dwelling fantail darters (Etheostoma flabellare) selected lower temperatures in winter (19.3°C) compared to pool dwelling johnny darters (E. nigrum; 22.0°C. A similar trend was evident in summer tests (fantail darters, 20.3°C; johnny darters, 22.9°C). Summer tested animals selected higher temperatures than winter tested animals maintained at the same acclimation temperature and photoperiod. When tested together in the same gradient, both species appeared not to thermoregulate, but tended to avoid each other. Critical thermal maxima (CTMax) did not differ between seasons for either species (fantail darters, 31.1°C winter, 31.3°C summer; johnny darters, 30.9°C winter, 30.5°C summer). Differences in the thermal responses of these darters correlated with differences in their respective habitats.     

PERMANENT GENETIC RESOURCES: Microsatellite DNA primers for the candy darter, Etheostoma osburni and variegate darter, Etheostoma variatum, and cross-species amplification in other darters (Percidae)

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.