Effects of egg composition and prey density on the larval growth and survival of lake whitefish (Coregonus clupeaformis Mitchill)

Eggs were collected from two stocks of lake whitefish, Coregonus clupeaformis , in Lakes Michigan and Huron to assess the effect of egg composition and prey density on larval growth and survival. Egg composition parameters including wet weight (mg egg⁻¹), dry weight (mg egg⁻¹), percent water, total caloric content (cal egg⁻¹), caloric density (cal egg⁻¹), percent lipid content, and total lipidcontent (mg egg⁻¹) were measured. Fish hatched from six parental females in each stock were fed one of four rations (0, 18, 24, 50 brine shrimp larva⁻¹ day⁻¹) after yolk sac absorption. Length at hatch, endogenous growth, exogenous growth, and survival were measured during a 42-day laboratory experiment. Length at hatch of larvae was positively related to egg caloric content ( r ²=0.780). Endogenous growth for lake whitefish larvae was positively related to percent lipid content ( r ²=0.896) and total egg lipid content ( r ²=0.876) of parental females. Exogenous growth and survival of larval lake whitefish was positively related to prey availability. Larval fish growth was accurately modelled ( r ²=0.973) as a function of prey abundance using a threshold-corrected hyperbolic equation. These results indicate that both egg composition and prey availability have the potential to influence the growth and survival dynamics of larval lake whitefish significantly.     

Population regulation in lake whitefish, Coregonus clupeaformis (Mitchill)

Mechanisms that might compensate for changes in mortality are well known but documentation of their operation and determination of their relative capacities is difficult. In this study the potential of lake whitefish to compensate for fishing mortality is quantified using a conventional fishery stock assessment model. The mechanisms examined are proportion of females, fecundity, survival of eggs and larvae, decrease in age at maturity and growth. The change in each of these parameters necessary to maintain constant recruitment with increasing fishing pressure is determined. Decrease in age of maturity and increased survival of larvae and eggs have the greatest potential for compensation. The total capacity for compensation appears large even when the mechanisms are considered alone. A review of several lake whitefish population studies indicates a large variation in the vital statistics. There is a close relation between total mortality and growth. There is also a close relation between growth and age at maturity. The analyses indicate that increased fishing results in increased growth which then results in a lower age at maturity. This feedback mechanism enables lake whitefish to respond to changes in environmental conditions and on average to maintain a birth rate in balance with mortality.     

Shoaling enhances cadmium avoidance by lake whitefish, Coregonus clupeaformis

Synopsis We compared the behavioural responses of solitary and shoaling lake whitefish, Coregonus clupeaformis, to Cd solutions, testing the hypothesis that fish are more responsive to contaminant gradients when in a shoal than when alone. The movements of individual fish were tracked in a countercurrent-type trough with clean water on one side, and water containing sequentially increasing Cd concentrations (0.2–125 g l^–1) on the other; the tracked fish was either alone, or within a shoal of four fish. We quantified a number of parameters derived from elements of locomotor behaviour and spatial selection (including the percent-time spent on the Cd-treated side) as potential indicators of response. Individuals in shoals responded to Cd more strongly and at lower concentrations than did solitary fish. While this was indicated by stronger avoidance of Cd, other response measures associated with escape behaviour illustrated this difference more clearly. Our results suggest that the social influence of shoaling enhances a fish's ability to respond to the presence of contaminants.     

Genetically based phenotype-environment association for swimming behavior in lake whitefish ecotypes (Coregonus clupeaformis Mitchill)

Abstract.— Studies of phenotype-environment associations in adaptive radiation have focused largely on morphological traits related to resource-based phenotypic differences. The genetic basis of adaptive behaviors implicated in population divergence remains poorly understood, as few studies have tested the hypothesis of behavioral phenotype-environment associations. We provide evidence of a phenotype-environment association for differential adaptive swimming behaviors through experiments conducted on dwarf, normal, and hybrid lake whitefish ( Coregonus clupeaformis ). Highly significant differences were observed for depth selection, directional changes, and burst swimming, implicating a genetic basis for these behaviors. Hybrid crosses revealed that depth selection is under additive genetic control, while dominance effects were suggested for directional changes and burst swimming. Estimates for the genetic basis of behavioral differentiation from an animal model were consistent with these observations. Comparative estimates of behavioral differentiation ( Q _ST) against neutral expectations ( F _ST) revealed pronounced departures from neutral expectations in all three behavioral phenotypes, consistent with the hypothesis that directional selection has driven the divergence of behavior in dwarf and normal lake whitefish ecotypes.     

Assessing feeding competition between lake whitefish Coregonus clupeaformis and round whitefish Prosopium cylindraceum

We collected lake whitefish Coregonus clupeaformis and round whitefish Prosopium cylindraceum from the main basin of Lake Huron and Georgian Bay in the Laurentian Great Lakes of North America to investigate details of diet and feeding tactics of these species in different seasons. Lake whitefish supports important commercial fisheries in Lake Huron and both species make use of habitats near the Bruce Nuclear Power Development, on the eastern shore of Lake Huron. Most fish of both species showed generalist feeding behavior, but some lake whitefish appeared to show specialist prey selection. The invasive spiny water flea Bythotrephes longimanus was an important component of the diet of both species. There was considerable dietary overlap between the whitefish species, but the ecological implications of these dietary overlaps are mitigated by the fact that dominant prey species differed in most seasons. We conclude that the potential for ecologically significant interactions between lake whitefish and round whitefish resulting from competition for similar benthic food resources in the main basin of Lake Huron is probably low [Current Zoology 56 (1): 109-117 2010].     

Experimental evidence for reduced hybrid viability between dwarf and normal ecotypes of lake whitefish (Coregonus clupeaformis Mitchill)

Forces driving the evolution of reproductive isolation among natural populations, as well as the mechanisms involved to maintain it, are still poorly understood. Because sympatric fish ecotypes mainly differ in phenotypic traits associated with occupying distinct trophic niches, it is generally believed that reproductive isolation is mainly driven by ecological divergent selection, excluding genome incompatibility as a basis for postmating isolation. We did cross experiments between dwarf and normal ecotypes of lake whitefish (Coregonus clupeaformis Mitchill) originating from distinct glacial refugia to test the hypothesis that their geographical isolation during the Pleistocene may have led to sufficient genetic divergence for the development of reproductive isolation between them before their secondary contact. Similar fertilization success in pure and hybrid crosses indicated the absence of gametic incompatibility between the two ecotypes. In contrast, daily embryonic mortality rates were 2.4–4.7 times higher in reciprocal hybrid crosses compared to pure crosses, which supports our working hypothesis. These results, along with previous morphological and population genetic studies, indicate that both genetic and ecological mechanisms may jointly act to promote speciation among northern freshwater fish ecotypes.     

Isolation,characterization and cross‐salmonid amplification of 31 microsatellite loci in the lake whitefish (Coregonus clupeaformis,Mitchill)

Coregonine fish represent the most successful evolutionary lineage of salmonids with Coregonus as the most speciose salmonid genus inhabiting numerous postglacial lakes across the northern hemisphere. We isolated and characterized 31 polymorphic microsatellite loci in Coregonus clupeaformis with an average number of 5.3 alleles per locus (range three to eight) and an overall expected heterozygosity of 0.74 ± 0.11. Two loci revealed significant linkage associations through analyses of mapping families. Six additional salmonid taxa assessed for cross‐species amplification revealed between 18 and 26 positive amplifications and between two and 12 polymorphic loci per species.     

A comparison of otolith and scale ages for western Labrador lake whitefish,Coregonus clupeaformis

Synopsis Scale ages underestimate otolith ages of western Labrador lake whitefish. Discrepancies occur from age 4 or 5 and can be considerable.     

Sex steroids in plasma of lake whitefish Coregonus clupeaformis during spawning in Lake Erie

Lake whitefish, Coregonus clupeaformis, were collected from the Western basin of Lake Erie during spawning. Free and conjugated (sulfated and glucuronidated) steroids including testosterone (T), 11-ketotestosterone (11-kT), estradiol-17beta (E2) and 17,20beta-dihydroxy-4-pregnen-3-one (17,20betaP) were measured in the plasma by radioimmunoassay. In males, the progression of spermiation was characterized by a significant decrease in plasma free steroids, whereas the levels of conjugated steroids remained similar and low, except for sulfated and glucuronidated testosterone. Plasma sex steroids did not correlate with the density or the motility of the spermatozoa. In females, the concentration of plasma T was significantly higher in preovulating than in ovulating females. The levels of E2 and 17,20betaP in ovulating lake whitefish exhibited large variations ranging from below detection limit to 0.9 ng ml(-1) and from 0.2 to 13 ng ml(-1), respectively. Analysis of conjugated steroids revealed high levels of glucuronidated and sulfated 17,20betaP and glucuronidated T in females ovulating in December. However, no significant differences in the proportion of the conjugated steroids were observed.     

Combining the analyses of introgressive hybridisation and linkage mapping to investigate the genetic architecture of population divergence in the lake whitefish (Coregonus clupeaformis, Mitchill)

Adaptation and reproductive isolation, the engines of biological diversity, are still elusive when discussing the genetic bases of speciation. Namely, the number of genes and magnitude of selection acting positively or negatively on genomic traits implicated in speciation is contentious. Here, we describe the first steps of an ongoing research program aimed at understanding the genetic bases of population divergence and reproductive isolation in the lake whitefish (Coregonus clupeaformis). A preliminary linkage map originating from a hybrid cross between dwarf and normal ecotypes is presented, whereby some of the segregating AFLP markers were found to be conserved among natural populations. Maximum-likelihood was used to estimate hybrid indices from non-diagnostic markers at 998 AFLP loci. This allowed identification of the most likely candidate loci that have been under the influence of selection during the natural hybridisation of whitefish originating from different glacial races. As some of these loci could be identified on the linkage map, the possibility that selection of traits in natural populations may eventually be correlated to specific chromosomal regions was demonstrated. The future prospects and potential of these approaches to elucidate the genetic bases of adaptation and reproductive isolation among sympatric ecotypes of lake whitefish is discussed.     

Accumulation, distribution and toxicology of dietary nickel in lake whitefish (Coregonus clupeaformis) and lake trout (Salvelinus namaycush)

An 18-day experiment was conducted to investigate the uptake and sublethal toxicity of dietary Ni in adult lake whitefish (LWF, Coregonus clupeaformis) and lake trout (LT, Salvelinus namaycush) fed diets containing 0, 1000 and 10000 microg Ni/g, prepared with and without brine shrimp. The results of this experiment were used to design an experiment of longer duration in which one of the fish species was selected and exposed to lower dietary Ni doses. In the present study feed refusal was observed in LT and LWF fed 10000 microg Ni/g, after three and 4-5 feedings, respectively. LT fed Ni-contaminated diets exhibited different patterns of Ni accumulation than LWF. Increased Ni concentrations in all LWF tissues, except the intestine, were associated with increased doses of Ni. Copper and Zn concentrations in kidney and liver of LWF were altered. Metallothionein concentrations in kidneys of LT fed 1000 microg Ni/g and 10000 microg Ni/g and LWF fed 10000 microg Ni/g and in livers of LWF fed 10000 microg Ni/g (diet without shrimp only) increased significantly. Increased lipid peroxide production in the plasma of LT fed 10000 microg Ni/g was observed. Blood glucose and electrolytes were affected by Ni exposure. Histopathological alterations were observed in kidneys of LWF fed low and high dose diets, livers of whitefish fed high dose diets, and intestines of LWF fed high dose diets and LT fed low and high dose diets. LT fed high dose diets exhibited significant decreases in weight.     

Real‐time PCR identification of lake whitefish Coregonus clupeaformis in the Laurentian Great Lakes

The purpose of this study was to develop a real‐time PCR assay to specifically identify lake whitefish Coregonus clupeaformis in larval fish assemblages based on a 122 bp amplicon from the mitochondrial genome. The efficiency of the reaction, as calculated from the standard curve, was 90·77% with the standard curve having an r² value of 0·998. Specificity of the assay provided single melt peak in a melt‐curve analysis and amplification of only the target species. The assay successfully identified target DNA in as low as 0·1% proportion of a DNA mixture. This assay was designed on the portable Smart Cycler II platform and can be used in both field and laboratory settings to successfully identify C. clupeaformis.     

Metazoan parasites of whitefish Coregonus clupeaformis (Mitchill) and cisco C. artedii Lesueur from Southern Indian Lake, Manitoba

Metazoan parasites of whitefish Coregonus clupeaformis and cisco C. artedii from Southern Indian Lake, Manitoba were studied to reveal: species composition, differences with host age, sex, and location and season of capture. Whitefish hosted 19 species, 18 of which were also in cisco with generally lower intensity levels. Parasites exhibited definite patterns of abundance with host age and season, the primary causes being dietary and behavioural. No differences in parasite abundance existed between host sexes. Ranking of cisco parasites was significantly different between two sampling sites while whitefish parasites did not differ. Whitefish and cisco from sites 40 miles (64 km) apart had significantly different abundances of Tetracotyle intermedia but not Triaenophorus crassus . An increase in the abundance of copepod-vectored cestodes with a concomitant decrease in abundance of amphipod-vectored parasites is predicted after flooding and diversion.     

Modifying effects of a cobble substrate on thermal environments and implications for embryonic development in lake whitefish (Coregonus clupeaformis)

A laboratory flume was constructed to examine substrate effects on aquatic development. The flume was designed as a once-through system with a submerged cobble-filled corebox. Lake whitefish (Coregonus clupeaformis) embryos and temperature probes were deployed at multiple sites within the cobble and in the open water channel. Embryos were incubated in the flume for two different experimental periods: one to examine substrate impacts during natural lake cooling (37 days: 5 December 2016 to 10 January 2017) and the second to investigate substrate effects while administering a twice weekly 1 h heat shock (51 days: 11 January to 2 March 2017). During incubation, no significant difference was found in the average temperature between locations; however, temperatures were more stable within the cobble. Following both incubation periods, embryos retrieved from the cobble were significantly smaller in both dry mass and body length by up to 20%. These results demonstrate differences between embryos submerged in a cobble substrate and in the open water column, highlighting the need to consider the physical influences from the incubation environment when assessing development effects as part of any scientific study or environmental assessment.     

BAC library construction, screening and clone sequencing of lake whitefish (Coregonus clupeaformis, Salmonidae) towards the elucidation of adaptive species divergence

Genomic DNA sequences and other genomic resources are essential towards the elucidation of the genomic bases of adaptive divergence and reproductive isolation. Here, we describe the construction, characterization and screening of a nonarrayed BAC library for lake whitefish (Coregonus clupeaformis). We then show how the combined use of BAC library screening and next-generation sequencing can lead to efficient full-length assembly of candidate genes. The lake whitefish BAC library consists of 181,050 clones derived from a single heterozygous fish. The mean insert size is 92 Kb, representing 5.2 haploid genome equivalents. Ten BAC clones were isolated following a quantitative real-time PCR screening approach that targeted five previously identified candidate genes. Sequencing of these clones on a 454 GS FLX system yielded 178,000 reads with a mean length of 358 bp, for a total of 63.8 Mb. De novo assembly and annotation then allowed retrieval of contigs corresponding to each candidate gene, which also contained up- and/or downstream noncoding sequences. These results suggest that the lake whitefish BAC library combined with next-generation sequencing technologies will be key resources to achieve a better understanding of both adaptive divergence and reproductive isolation in lake whitefish species pairs as well as salmonid evolution in general.     

Contrasting patterns of mitochondrial DNA and microsatellite introgressive hybridization between lineages of lake whitefish (Coregonus clupeaformis); relevance for speciation

We performed a combined analysis of mitochondrial DNA (mtDNA) and microsatellite loci among lake whitefish (Coregonus clupeaformis) populations in order to assess the levels of congruence between both types of markers in defining patterns of genetic structuring, introgressive hybridization and inferring population origins in the hybrid zone of the St. John River basin. A second objective was to test the hypothesis that secondary contact between glacial lineages always resulted in the occurrence of sympatric dwarf and normal whitefish ecotypes. Fish were sampled from 35 populations and polymorphism was screened at mtDNA and six microsatellite loci for a total of 688 and 763 whitefish, respectively. Four lakes harbouring a single whitefish population of normal ecotype admixed with mtDNA haplotypes of different lineages were found. This confirmed that secondary contact between whitefish evolutionary lineages did not always result in the persistence of reproductively isolated ecotypes. Microsatellites further supported the definition of distinct glacial lineages by identifying lineage-specific allelic size groups. They also further supported the hypothesis that ecotypes originated from either a single founding lineage (sympatric divergence) or following secondary contacts between lineages (allopatric divergence), depending on the lake. In general, however, the pattern of population differentiation and introgressive hybridization observed at microsatellites was in sharp contrast with that depicted by mtDNA variation. Both factorial correspondence analysis and analysis of admixture proportion revealed a much more pronounced pattern of introgressive hybridization than depicted by mtDNA analyses. Variable levels of introgression indicated that environmental differences may be as important as the historical contingency of secondary contact in explaining the persistence of sympatric ecotypes and the differential pattern of introgressive hybridization among lakes. Whitefish populations from the St. John River basin hybrid zone represent a rare illustration of a continuum of both morphological and genetic differentiation within a given taxon, spanning from complete introgression to possibly complete reproductive isolation, depending on lakes. Thus, each lake may be viewed as a different temporal snapshot taken throughout the gradual process of speciation.     

Collapse of Lake Whitefish Coregonus clupeaformis (Mitchill, 1818) species pair in Como Lake,Ontario

Lake Whitefish (Coregonus clupeaformis) species pairs exist in a small number of northern temperate lakes in North America. Sympatric populations of dwarf and normal‐sized whitefish represent a continuum of morphological and genetic divergence among these lakes. In this study, we used morphological and age data to assess the status of the Lake Whitefish species pair in Como Lake, Ontario (Canada). Whitefish were collected with gill‐nets from spawning shoals during the fall of 2012 and 2014 and compared with archived (circa 1989) dwarf and normal‐sized specimens. Our study indicates that the two previously collected forms have been replaced by a single form with a different morphology and age structure. In comparison to archived specimens, contemporary Lake Whitefish were larger, deeper‐bodied and older; with more gill rakers, lateral line scales, smaller eyes and a wider interorbital width. We hypothesize that the recent invasion of Como Lake by Spiny Waterflea (Bythotrephes longimanus) has affected whitefish growth rates and changed the ecosystem conditions that previously maintained the species pair.