Allopaternal care in the tessellated darter,Etheostoma olmstedi (Pisces: Percidae)

Synopsis Large male tessellated darters, Etheostoma olmstedi, defend flat-bottomed rocks, the undersides of which serve as spawning substrates. Because females attach eggs directly onto bare stone, a spawning bout necessarily decreases the surface area available for further breeding at a nest. In apparent response to the female preference of ovipositing on rocks with the most bare surface, fathers abandon nests with their attached offspring to search for rocks with more uncovered surface. Egg abandonment also results after roving fathers intercept and spawn with ripe females and remain at a different rock to care for a new clutch. In essence, large males appear to maximize the number of eggs they fertilize by sequentially monopolizing breeding patches that are temporarily most desirable to females. The costs to a father of abandoning his eggs appear to be low for two reasons: (1) non-territorial males, created by a scarcity of suitable rocks, readily occupy vacant nests, and clean and aerate the abandoned eggs; and (2) after water hardening, eggs are less vulnerable to cannibalism. The maintenance of eggs by nonparental males may be both incidental to scrubbing the ceiling for their own breeding and adaptive in that viable eggs stimulate females to spawn; neither would appear to involve parental investment costs. Thus, allopaternal care in the tessellated darter may have evolved because it is performed in the selfish pursuit of spawning opportunities while entailing little, if any, of the costs normally subsumed within parental investment.     

Phylogeography of the greenside darter complex, Etheostoma blennioides (Teleostomi: Percidae): a wide-ranging polytypic taxon

The greenside darter, Etheostoma blennioides (Teleostomi: Percidae), is a wide-ranging polytypic taxon that occurs throughout eastern North America. A previous morphological study recognized four subspecies (blennioides, newmanii, gutselli, and pholidotum), several morphological races, and three zones of morphological intergradation. We generated complete cytochrome b (1140bp) sequence data for 51 individuals from across the range of the greenside darter inclusive of all of the currently recognized taxa to assess genetic variation and taxonomic boundaries. Both maximum parsimony and mixed model Bayesian analyses resulted in two strongly supported deeply divergent clades including (1) a Tennessee River drainage clade, and (2) an Ohio River and Great Lakes basins, Interior Highlands, and Atlantic slope clade. Etheostoma blennius, a closely related congener, nested within the Tennessee River clade of E. blennioides, rendering the complex paraphyletic. Test of alternative topologies failed to support the current taxonomic designations. The inclusion of nuclear sequence data from intron 1 of the S7 ribosomal protein (523bp) from a subset of the populations was included to independently test whether the currently recognized taxa conform to distinct evolutionary lineages and also to clarify potential issues associated with ancestral hybridization. Although the nuclear data was less variable than the mitochondrial data, the monophyly of several of the subspecies could not be rejected.     

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.     

Diets of four sympatric species of Etheostoma (Pisces: Percidae) from southern Indiana: interspecific and intraspecific multiple comparisons

Synopsis The diets of four species of Etheostoma (E. spectabile, E. caeruleum, E. flabellare and E. nigrum) were investigated from ten Order 3 streams in the White River drainage of southern Indiana. All species fed mainly on insect larvae, primarily chironomids, ephemeropterans and plecopterans. Dietary proportions, as frequency of occurrence, were compared using cluster analysis based on matrices of values of Spearman rank correlation, Schoener index, Jaccard association, and Pearson's r. Certain species and, in some cases, stream pairs within species clustered closely in all analyses or in three of the four. Most clusters do not show closer intraspecific than interspecific similarity and specific streams had more influence than species in forming the logic of some clusters. It can be concluded that these fishes are opportunistic predators.     

Characterization of eight polymorphic microsatellite DNA markers for the greenside darter,Etheostoma blennioides (Percidae)

The greenside darter, Etheostoma blennioides is a small benthic fish found in fast‐flowing streams in eastern North America. In Canada, this species is native to three, and introduced into one, Great Lakes tributaries in southwestern Ontario. It is currently listed as a species of Special Concern. To characterize population genetic structure and diversity in the Canadian populations of greenside darter, eight polymorphic microsatellite markers were developed for the species. The polymerase chain reaction primers were tested between 32 and 60 individuals from the Sydenham River and yielded a high number of alleles (four to 42 per locus), and observed heterozygosities ranging from 0.14 to 0.82.     

Microsatellite DNA primers for the endangered vermilion darter,Etheostoma chermocki,and cross‐species amplification in other darters (Percidae: Etheostoma)

The endangered vermilion darter (Etheostoma chermocki) is endemic to the Black Warrior River system in the Mobile Basin in Alabama. Restoration and conservation of this species require an understanding of its population genetic structure, which can be characterized using microsatellite DNA. Nine microsatellite loci were developed; eight loci were polymorphic. Although observed heterozygosity was lower than expected heterozygosity in most polymorphic loci, only one locus showed significant deviation from Hardy–Weinberg equilibrium. These nine markers were tested in an additional 24 species of Etheostoma and appear to have sufficient allelic variation to be useful in studies of population genetic structure.     

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.     

Spawning and nesting behavior of the waccamaw darter,Etheostoma perlongum

Synopsis The spawning and nesting behavior ofEtheostoma (Boleosoma) perlongum was investigated in the field and laboratory. Sexual dimorphism is highly developed in such features as genital papillae, first dorsal and paired fins, and nuptial coloration. A reproductive migration from mid-lake to shore occurs in the spring: males precede females to select nest sites under submerged sticks and other debris. The male excavates a depression beneath the submerged object. Gonad analysis indicates a single spawning season extending from March through June. Nests were found from late April to mid-June and were guarded by a single male for periods of 13 to 36 days. Males initiate courtship by lateral display, lead the female to the nest site and show the nest by inverting. The female responds by tail up, tail wag and circle; males also tail wag and circle. Spawning pairs invert, usually in unison, and orient head to head or, less often, head to tail. The female deposits eggs while holding her body in a weak S or J shape with the caudal peduncle held away from the spawning substrate while vibrating.     

Seasonality,aestivation and the life history of the salamanderfish Lepidogalaxias salamandroides (Pisces: Lepidogalaxiidae)

Synopsis Changes in the age/length structure of Lepidogalaxias salamandroides populations in temporary waters of south-western Australia were monitored over a 51 month period by regular field sampling. Each year the study area experienced a summer drought of approximately five months duration. During the drought period Lepidogalaxias burrows into the mud and aestivates and substantial mortality occurs during this period. Body lipid reserves decrease during this period suggesting that they are the main energy source used during aestivation. The amount of lipid remaining after aestivation appears to be important in determining when female fish reproduce. Males die after reproduction and achieve a maximum age of only about 12 months. Some females also reproduce and die at this age while others defer reproduction for another year. Fecundity is much higher in these older, larger fish: they must however, survive two drought periods before they reproduce. The evolution of the life history style shown by Lepidogalaxias salamandroides is discussed with respect to the well defined seasonality of the region and it is suggested that selection for reproduction at an early age is balanced by the constraints of offspring survival during the aestivation period.     

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.     

Clutch and egg size variation in the banded darter, Etheostoma zonale, from three sites in Arkansas

Synopsis We studied variation in clutch and egg size of the banded darter, Etheostoma zonale, using museum specimens representing three geographically separated populations from central and northern Arkansas. We found significant differences in standard length-adjusted mean clutch size and mean ovum mass. Clutch size varied by more than three hundred percent between sites, and egg mass by approximately twenty percent, demonstrating that comparative studies of interspecific variation in life history based upon a single population for each species may be misleading.     

Life history variation among female surfperches (Perciformes: Embiotocidae)

Synopsis Life history variation within the family Embiotocidae is extensive and involves differences in age of first reproduction, fecundity schedules, growth rates, longevity and size of young. Based on maximum reported body lengths, there are three distinct size groups among the family's 23 species. Small species do not exceed 215 mm TL, medium-size species attain 275 to 335 mm TL, and the large species attain 380 to 470 mm TL. The longevity oh surfperches varies from two to ten years, growth is indeterminate, and females of the medium-and large-size groups may delay first reproduction beyond age one. With one exception, all species show increasing length-specific fecundities. The life history characteristics of females differ among the three size groups. Relative to smaller species, the largest species have moderately high fecundity, delayed maturity and long life. Medium-size species have low fecundity, may delay maturity for 1 to 3 years and have intermediate life spans. Small species have generally higher, but variable, fecundity, do not delay maturity, and are short lived. Among the small North American species, the trend in fecundity varies inversely with environmental predictability. Fecundity is highest in the species which occupies highly seasonal freshwater environments. Coastal species produce moderately large broods and species which occupy stable deep water environments produce the smallest broods.     

Observations on the life history of the belted sandfish,Serranus subligarius (Serranidae)

Synopsis The biology of Serranus subligarius, a typical inhabitant of low limestone reef outcroppings and artificial reefs from North Carolina to Florida and in the northern Gulf of Mexico, was studied on inshore jetties in the northeastern Gulf of Mexico from February 1974 to April 1975. The length-weight relationship was expressed by the equation W = 0.000006L^3.431. Growth was greatest in the first year (43.5 mm SL) and declined in the remaining three years of life (17.3, 10.5, and 7.1, respectively). In individuals larger than 40 mm SL, gammaridean and caprellid amphipods were the most numerous food items but crabs, shrimp, and fishes were the most important food items by weight. In individuals less than 40 mm SL, copepods were the most numerous food item, but gammaridean amphipods and shrimp were the most important items by weight. Two incidents of cannibalism were recorded. The ovotestis of the belted sandfish is composed of two lobes which are broadly united posteriorly, with the testicular tissue restricted to a broad band encircling the posterior region of the gonad. Age-group 1 fish are functional synchronous hermaphrodites. Testicular tissue matures in the fall or winter of a fish's first year (all specimens examined histologically, including those from the nonspawning season, contained flagellated spermatozoa) while ovarian tissue matures in the following spring. Spawning season in 1974 extended from late April or early May to September. Crossfertilization is the usual reproductive mode, although individuals are capable of selffertilization.     

Antimicrobial egg cleaning by the fringed darter (Perciformes: Percidae: Etheostoma crossopterum): implications of a novel component of parental care in fishes

Broad-spectrum antimicrobial compounds have recently been identified in the epidermal mucus of fishes and probably serve as a first line of defence against microbial pathogens. Because of the ubiquitous nature of fungi and bacteria in aquatic systems, defence against these pathogens should be required throughout the lifespan of fishes, including the egg stage. We conducted experiments on Etheostoma crossopterum (Percidae: Catonotus), the fringed darter, to determine if the presence of a guarding male inhibits microbial colonization of eggs. Based on results from a combination of in-stream experiments, in vitro microbial assays, and morphological characteristics and behaviour of breeding males, we propose that antimicrobial egg cleaning by the guarding male is an effective component of parental care in these fish. Although innate antimicrobial compounds have been identified in a variety of organisms ranging from insects to vertebrates, integration of these compounds into a species's reproductive life history has been identified only in a small number of insect species. The results from this study not only indicate that E. crossopterum males provide a novel form of vertebrate parental care, but also have implications regarding the evolution of parental care in fishes and transitional evolutionary stages from no parental care to male parental care.     

Systematics of the subgenus Oligocephalus (Teleostei: Percidae: Etheostoma) with complete subgeneric sampling of the genus Etheostoma

The genus Etheostoma is the most diverse clade of freshwater fishes in North America. While studies have been performed with complete sampling of a single subgenus, none have included representatives of all remaining subgenera. The subgenus Oligocephalus is the largest, consisting of 25-27 species in four species groups, and its monophyly has never been clearly demonstrated. The monophyly of this subgenus and its constituent groups was tested using parsimony and Bayesian analyses of ND2 (mtDNA) and the first intron of S7 (nDNA) with complete species sampling from Oligocephalus and complete subgeneric sampling from Etheostoma. Although the subgenus Oligocephalus was not recovered as a monophyletic group in any analyses, monophyletic E. whipplei, Southwestern Darter, and E. spectabile (in part) species groups were recovered in all analyses. All analyses agree that E. okaloosae and both subspecies of E. hopkinsi are not closely related to other members of the subgenus Oligocephalus. E. exile is, however, presenting the strongest evidence yet that recognition of the subgenus Boleichthys is unwarranted.