Analysis of the effect of transplantation on morphometric and meristic characters in lake populations of the rainbow smelt,Osmerus mordax (Mitchill)

Synopsis Results of experimental transfer of rainbow smelt into lakes reclaimed by rotenone around 1960 in Maine were originally interpreted to cast doubt on the previously widely accepted hypothesis that there were two hereditarily different forms of rainbow smelt, one large and one small. Study of more recent data from some of the transplanted populations and reanalysis of the original data suggests different conclusions. The initial effect of introductions into a reclaimed lake may be accelerated and/or more prolonged growth which exceeds even interlake differences. This initial phase, however, is followed by a second phase, when the population reaches equilibrium and these effects subside. Data from phase two of the Little Concord-Shagg and Cold Stream-Coleback Lake transfers showed that the growth characteristics of the transplanted populations returned to those of the parental populations. Large differences in growth patterns were thus found only in the initial phase of the introduction. Meristic characters were little affected by transplanting.Analysis of large specimens derived from a postulated second unofficial introduction into Coleback Lake showed that they also differed significantly, having both higher gill raker and vertebral counts than the smaller smelt. This was of interest as smelt vertebral and gill raker counts usually are inversely related; hence we do not equate these for the moment with the large form of smelt known elsewhere.It is concluded that the initial interpretation of transfer experiments be delayed until conditions approaching equilibrium can be expected to exist. Further, our analysis of more recent lake transfer data has shown nothing to refute the hypothesis that there are at least two hereditarily different forms of smelt.     

The New Zealand common smelt: biology and ecology

The common smelt is one of the most widespread indigenous freshwater fishes in New Zealand. One other member of the family Retropinnidae, Stokellia anisodon (Stokell), is present but is confined to a small region of the South Island. There are many diadromous as well as river and lake resident populations, the latter, sometimes a result of introductions to serve as forage fish for trout. Diadromous smelt spawn during austral autumn–winter on sand bars of lower riverine reaches. Larval stages inhabit coastal marine waters, and the postlarvae to immature stages re‐enter rivers and some lowland lakes. Diadromous smelt are distinguished from lowland lake resident forms by high vertebral but low gill raker numbers and larger size and from those present in some isolated waters, by high vertebral numbers alone. Lake or reservoir resident smelt usually spawn in austral spring–summer on sandy shallows at stream mouths or along shorelines. Verified smelt ages (otolith analyses) indicate that in some populations most smelt mature and spawn after c. 1 year. Adult smelt feed on a spectrum of primarily invertebrate animals ranging from small zooplankters to insects and occasionally small fishes. Smelt are a major prey for both brown trout and rainbow trout. Adult smelt are a minor food for the Maori people. As postlarvae they are a component of a few 'whitebait' fisheries. Most smelt populations are increasingly affected by environmental changes induced by human activities. Although many studies have examined problems affecting smelt, further effort is required, along with more basic research.     

Effects of elevated turbidity on shallow lake fish communities

Synopsis We compared the fish communities of two shallow lakes in the lower Waikato River basin, North Island, New Zealand, to determine the effects of elevated suspended solids (SS) and collapse of submerged macrophytes. Lake Waahi was turbid (20–40 g m^-3 SS) and devoid of submerged macrophytes whereas Lake Whangape was clearer (5 g m^-3 SS) and dominated by submerged macrophytes. The lakes had similar fish species richness and had nine major species in common; representing eight families including Anguillidae, Retropinnidae, Galaxiidae, Eleotridae, Mugilidae, Ictaluridae, Poeciliidae, and Cyprinidae (two species). The only major fish that was absent from Lake Waahi was a lacustrine form of the common smelt, Retropinna retropinna, which disappeared after the lake became turbid in the late 1970s. CPUE, condition, and size of most species in Lake Waahi were similar to, or greater than, those in Lake Whangape. Lake Whangape clearly exceeded Lake Waahi only for CPUE of two species. Within Lake Whangape two species displayed significantly greater condition, and one species greater size, in a turbid arm of the lake than in the main basin. Apart from lacustrine Retropinna retropinna, the fish in these lakes appear well adapted to cope with, or to avoid, the direct toxic effects of suspended and settleable solids on sensitive early developmental stages. In Lake Waahi loss of cover and food provided by submerged macrophytes appears to have been compensated for by increased turbidity and an associated increase in the biomass of the mysid, Tenagomysis chiltoni (a major prey item).     

Validation of Alburnus timarensis from the Lake Van basin,eastern Anatolia (Teleostei: Cyprinidae)

The Lake Van basin located in eastern Anatolia is inhabited by two species of Alburnus: Alburnus tarichi is a migratory species foraging in the lake and spawning in all tributaries, while A. timarensis is resident in streams and is known only from one tributary of the lake, the Karasu. It could be shown that A. timarensis is a valid species, which is rediagnosed here. Both species are syntopic during the spawning season, but are well distinguished by size, gill raker and lateral line scale counts.     

Coexistence and origin of trophic ecotypes of pygmy whitefish,Prosopium coulterii,in a south‐western Alaskan lake

Ecologically, morphologically and genetically distinct populations within single taxa often coexist in postglacial lakes and have provided important model systems with which to investigate ecological and evolutionary processes such as niche partitioning and ecological speciation. Within the Salmonidae, these species complexes have been well studied, particularly within the Coregonus clupeaformis–C. laveratus (lake and European whitefish, respectively) group, but the phenomenon has been less well documented in the other whitefish genera, Prosopium and Stenodus. Here, we examined the morphology, feeding biology and genetic structure of three putative forms of the pygmy whitefish, Prosopium coulterii (Eigenmann & Eigenmann, 1892), first reported from Chignik Lake, south‐western Alaska, over 40 years ago. Field collections and morphological analyses resolved a shallow water (< 5 m depth) low gill raker count form (< 15 first arch gill rakers), a deepwater (> 30 m), low gill raker form and a deepwater, high gill raker count (> 15 gill rakers) form. The two low gill raker count forms fed almost exclusively on benthic invertebrates (mostly chironomids), while the deepwater, high gill raker count form fed almost exclusively on zooplankton; differences in diet were also reflected in differences both in δ¹³C and δ¹⁵N stable isotopes. All three forms were characterized by the same major mitochondrial DNA clade that has been associated with persistence in, and postglacial dispersal from, a Beringian glacial refugium. Analysis of variation at nine microsatellite DNA loci indicated low, but significant differentiation among forms, especially between the two low gill raker count forms and the high gill raker count form. The extent of differentiation along phenotypic (considerable) and genetic (subtle) axes among the Chignik Lake forms is similar to that found among distinct taxa of Prosopium found in pre‐glacial Bear Lake (Utah–Idaho, USA) which is probably at least ten times older than Chignik Lake. Our analyses illustrate the potential for the postglacial differentiation in traits subject to divergent natural selection across variable environments.     

Molecular genetic evidence for reproductive isolation between sympatric populations of smelt Osmerus in Lake Utopia, south-western New Brunswick, Canada

Two morphologically and ecologically distinct forms of smelt, Osmerus, reside sympatrically in Lake Utopia, south-western New Brunswick, Canada. The ‘normal-sized’ form matures at greater than 200 mm standard length, averages about 31–33 gill rakers, and spawns in lake outlets. By contrast, the ‘dwarf-sized’ form matures at less than 150 mm standard length, averages 34–36 gill rakers, and spawns in small streams 3–5 weeks later than the normal form. We tested whether these sympatric forms represented ecological polymorphism within a single population or two reproductively isolated demes by assaying variation within and between forms by mitochondrial DNA (mtDNA) restriction site and nuclear minisatellite DNA analyses. Analysis of smelt mtDNA with twelve restriction enzymes resolved ten composite genotypes (differing by an average 0.27% sequence divergence) which differed markedly in frequency between the forms. Net percentage sequence divergence between the forms was O.l6%. A Wagner parsimony/ bootstrapping analysis of the restriction site presence/absence matrix, however, suggested that there were no significant distinctions between dwarf and normal smelt based on the phylogeny of composite genotypes. Hybridization studies of genomic DNA digests with a minisatellite probe indicated both that nuclear restriction fragment differentiation and the frequency of specific fragments differed significantly between the forms. Significant genetic differentiation between the sympatric forms demonstrates that they are distinct gene pools and reproductively isolated. Our molecular evidence for reproductive isolation between dwarf and normal smelt in Lake Utopia, coupled with the persistent morphological and ecological differentiation between them, argues strongly that they are behaving as distinct species. The Lake Utopia Osmerus populations provide further illustration of the potential for rapid differentiation to the level of biological species in postglacial environments.     

Going,going, gone: evidence for loss of an endemic species pair of threespine sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>) with implications for protection under species-at-risk legislation

Genomic extinction occurs when the unique combination of genetic traits that characterize distinct phenotypes are eliminated by introgressive hybridization even if population size is greater than zero. Benthic and limnetic threespine sticklebacks (Gasterosteus aculeatus) constitute reproductively isolated undescribed biological species that have evolved independently in several lakes in southwestern British Columbia, Canada (known as “species pairs” in each lake). Here we investigated whether the two species that comprise the pair from Enos Lake, southeastern Vancouver Island, remain as two distinct gene pools. Multi-season samples (>1200 fish) obtained over two years from throughout the lake and assayed for variation in morphological traits characteristic of the two species (i.e., body depth, dorsal spine count, gill raker counts) and at 12 microsatellite DNA loci consistently indicated the existence of only a single group of sticklebacks. There was no consistent evidence of two groups in any morphological trait, and mean gill raker counts were consistently intermediate (20–21) to those of known benthics (~18) and limnetics (~24) which together comprised strikingly bimodal distributions in historical samples. Genetic analyses employing model-based clustering also consistently indicated the presence of only a single genetic group of sticklebacks. Compared to historical samples and to benthics and limnetics from other lakes, no Enos Lake fish could be identified confidently as a pure benthic or limnetic. Our results provide the strongest evidence yet that the Enos Lake sticklebacks now consist of a single morphological and genetic population of sticklebacks, that the unique combination of genetic and morphological traits that characterized benthic and limnetic sticklebacks no longer exist, and that their current status under Canada’s Species-at Risk Act as Endangered should be re-evaluated.     

Resident spawning of Japanese smelt,Hypomesus nipponensis,along gravel shorelines in Lake Nojiri,Central Japan

In Lake Nojiri, the Japanese smelt, Hypomesus nipponensis McAllister, 1963, population has been thought to be maintained by artificial spawning, and stocking efforts as natural spawning in in-flowing streams is unlikely due to the lack of inflowing stream habitat. In this study, novel resident Japanese smelt spawning was observed along the lakeshore in shallow areas with clean gravel and flow. Spawning occurred at night during early March to mid-April and eggs, confirmed on gravel substrate, progressed to the eyed state in 5–6 weeks. Although natural spawning was observed, the ratio of eggs that progressed to the eyed stage was minimal, possibly due to thick algae cover in areas with low flow velocity. Therefore, resident Japanese smelt spawning along the lake shoreline is physically possible and potentially contributes to the overall population in Lake Nojiri, but in-lake spawning alone presumably is not a biologically viable method for population sustainability due to low survival rates and egg density at present. Environmental improvements such as construction of fish ladders to suitable in-flowing spawning habitat, introduction of clean gravel to shorelines, and water level management adjusted to the smelt spawning run would contribute to higher recruitment by natural smelt reproduction, and consequently enhance the smelt production in Lake Nojiri.

     

Patterns of morphological changes and hybridization between sympatric whitefish morphs (Coregonus spp.) in a Swiss lake: a role for eutrophication?

Whitefish, genus Coregonus, show exceptional levels of phenotypic diversity with sympatric morphs occurring in numerous postglacial lakes in the northern hemisphere. Here, we studied the effects of human‐induced eutrophication on sympatric whitefish morphs in the Swiss lake, Lake Thun. In particular, we addressed the questions whether eutrophication (i) induced hybridization between two ecologically divergent summer‐spawning morphs through a loss of environmental heterogeneity, and (ii) induced rapid adaptive morphological changes through changes in the food web structure. Genetic analysis based on 11 microsatellite loci of 282 spawners revealed that the pelagic and the benthic morph represent highly distinct gene pools occurring at different relative proportions on all seven known spawning sites. Gill raker counts, a highly heritable trait, showed nearly discrete distributions for the two morphs. Multilocus genotypes characteristic of the pelagic morph had more gill rakers than genotypes characteristic of benthic morph. Using Bayesian methods, we found indications of recent but limited introgressive hybridization. Comparisons with historical gill raker data yielded median evolutionary rates of 0.24 haldanes and median selection intensities of 0.27 for this trait in both morphs for 1948–2004 suggesting rapid evolution through directional selection at this trait. However, phenotypic plasticity as an alternative explanation for this phenotypic change cannot be discarded. We hypothesize that both the temporal shifts in mean gill raker counts and the recent hybridization reflect responses to changes in the trophic state of the lake induced by pollution in the 1960s, which created novel selection pressures with respect to feeding niches and spawning site preferences.     

Contemporary trait change in a classic ecological experiment: rapid decrease in alewife gill‐raker spacing following introduction to an inland lake

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Analyses of growth processes of pond smelt, Hypomesus nipponensis, in Lake Ogawara, Japan, through the use of daily otolith increments

It has been reported that pond smelt, Hypomesus nipponensis, in Lake Ogawara, Japan, appear in small and large size groups during spawning despite being an annual fish. It is hypothesized that pond smelt have a bimodal life history, anadromous for large size groups and resident in the lake for small size groups. We calculated the body length and growth rate of the small and large size groups through the use of daily otolith increments and compared growth parameters between groups. In addition, the growth processes of resident fish in Lake Ogawara and anadromous fish in the adjacent sea were examined and compared with small and large size groups, respectively. We found that the two size groups diverged after 40–50 days from hatching with significant size groups present after 50 days. Through otolith increment analysis the growth processes of the small and large size groups were correlated with resident and anadromous forms respectively. These results revealed the utility of otolith increment analyses to clarify population structure of this species.     

Adaptive divergence between lake and stream populations of an East African cichlid fish

Divergent natural selection acting in different habitats may build up barriers to gene flow and initiate speciation. This speciation continuum can range from weak or no divergence to strong genetic differentiation between populations. Here, we focus on the early phases of adaptive divergence in the East African cichlid fish Astatotilapia burtoni, which occurs in both Lake Tanganyika (LT) and inflowing rivers. We first assessed the population structure and morphological differences in A. burtoni from southern LT. We then focused on four lake–stream systems and quantified body shape, ecologically relevant traits (gill raker and lower pharyngeal jaw) as well as stomach contents. Our study revealed the presence of several divergent lake–stream populations that rest at different stages of the speciation continuum, but show the same morphological and ecological trajectories along the lake–stream gradient. Lake fish have higher bodies, a more superior mouth position, longer gill rakers and more slender pharyngeal jaws, and they show a plant/algae and zooplankton‐biased diet, whereas stream fish feed more on snails, insects and plant seeds. A test for reproductive isolation between closely related lake and stream populations did not detect population‐assortative mating. Analyses of F1 offspring reared under common garden conditions indicate that the detected differences in body shape and gill raker length do not constitute pure plastic responses to different environmental conditions, but also have a genetic basis. Taken together, the A. burtoni lake–stream system constitutes a new model to study the factors that enhance and constrain progress towards speciation in cichlid fishes.     

Disappearance of koaro,Galaxias brevipinnis,from Lake Rotopounamu,New Zealand,following the introduction of smelt,Retropinna retropinna

Synopsis Koaro, Galaxias brevipinnis, were once the only fish present in Lake Rotopounamu but, after a comprehensive survey in 1990, none were found in the lake or its tributary streams. Introduced native fish, specifically smelt, Retropinna retropinna, and the common bully, Gobiomorphus cotidianus, now occur in this lake. As koaro co-exist with bullies in other lakes, but have declined in landlocked lakes containing smelt, the disappearance of koaro in Lake Rotopounamu is attributed to the introduction of smelt alone. Interspecific competition for food between 0 + year old koaro and smelt, combined with predation by 2 + year old smelt on koaro larvae, are thought to be responsible. Such a mechanism would be consistent with theoretical predictions of predator-prey regulation systems within same chain food webs. Rainbow trout, Oncorhynchus mykiss, which were introduced into a number of local lakes before smelt, and which preyed on the koaro, have been blamed for the decline of the koaro populations. However, the disappearance of koaro in Lake Rotopounamu shows that smelt can reduce koaro populations independently of trout predation.     

Ecological niche specialization inferred from morphological variation and otolith strontium of Arctic charr Salvelinus alpinus L. found within open lake systems of southern Baffin Island, Nunavut, Canada

The presence of two morphotypes of Arctic charr Salvelinus alpinus was confirmed via morphological variation and otolith strontium (Sr) within three open-lake systems of southern Baffin Island, Nunavut, Canada: Qinngu (LH001), Iqalugaarjuit Lake (PG082) and Qasigiat (PG015). Analysis of otolith Sr indicates that a component of each S. alpinus population within lakes LH001 and PG082 is migratory (large–maturing S. alpinus ), whereas another component is lake-resident (small–maturing S. alpinus ). Alternatively, small and large maturing S. alpinus may both inhabit tidal habitats during their lifetime in lake PG015. Three morphological characters were identified by principal factor analysis (PFA) as characters that were different between maturity groups for all lakes studied: eye diameter, pectoral fin length and pelvic fin length. As well, upper jaw length (LH001 and PG082) and fork depth (PG015) were identified in PFA as traits that differed between morphs. Univariate tests of morphological characters identified by PFA demonstrated maturity group differences with the exception of eye diameter in Lake PG015 and upper jaw length and pelvic fin length in lake LH001. No difference was found in the MANOVA test of upper and lower gill raker number between small–maturing and undeveloped fish within all lakes studied. Clear morphological variation observed between small–maturing and undeveloped fish in all three lakes of the study suggests ecological niche separation between morphotypes. This is the first documented case of lake-resident S. alpinus use of the tidal habitat in the presence of a migratory large–maturing morphotype.     

Meristic and morphometric variation in the silver perch, Leiopotherapon plumbeus (Kner, 1864), from three lakes in the Philippines

A number of researchers have applied multivariate methods to elucidate the population structure of fishes. In this study, we also used multivariate techniques to examine meristic and morphological variations in the silver perch, Leiopotherapon plumbeus (Kner, 1864), from three Philippine lakes, namely, Laguna de Bay, Sampaloc Lake, and Taal Lake. We also aimed to determine whether or not there are meristic and morphological differences among the populations of silver perch from the three lakes, considering that this fish species was introduced from Laguna de Bay into Sampaloc Lake and Taal Lake in the late 1950s and early 1970s, respectively. A total of 710 specimens from four different sites were used in the study: 155 each from Binangonan and Tanay areas of Laguna de Bay and 200 each from Sampaloc Lake and Taal Lake. Based on analysis of variance, eight meristic and 26 transformed morphometric characters were selected for subsequent analyses. Nineteen (19) principal components extracted from the 34 significant variables accounted for 82.3% of the variation in the original variables. Factor analysis using varimax rotation produced four factors: factor 1 was dominated by fin measurements while the highest loadings for factor 2 were gill raker counts. Factors 3 and 4 were dominated by various body and head measurements. Cluster analysis showed specimens from Sampaloc Lake and Taal Lake in one group, while majority of the specimens from Binangonan and Tanay are in another cluster. This suggests a closer morphological similarity between specimens from Sampaloc and Taal. Discriminant analysis gave relatively high correct classification rates (76.13–95.50%). Lower gill raker count was the most discriminating variable. Since both the silver perch from Sampaloc Lake and Taal Lake were introduced from Laguna de Bay, the observed clustering and morphological variation could be attributed to similarities and differences in the lake environments. Laguna de Bay is a shallow eutrophic lake, while Sampaloc Lake and Taal Lake are deep lakes. Further studies, however, are needed to determine which of the myriad of biological and/or physico–chemical factors might have the greatest influence on the observed morphological divergence between the source population and transplanted populations that we found in our study.     

Isolation between sympatric anadromous and resident threespine stickleback species in Mud Lake,Alaska

A sympatric pair of anadromous and resident freshwater threespine stickleback species (Gasterosteus aculeatus species complex) occurs in Mud Lake in the Matanuska-Susitna Valley, Alaska. The two forms differ in an array of morphological traits, including traits associated with predator defense (e.g., spine lengths) and trophic ecology (e.g., number of gill rakers). Mud Lake is only the third lake reported to have anadromous stickleback (which have a complete row of lateral plates) coexisting with low-plated resident stickleback in the absence of intermediate partially plated fish. Microhabitat and seasonal isolation appear to contribute to reproductive isolation between the two forms.     

Variation in number of vertebrae and gill rakers of sockeye salmon,Oncorhynchus nerka,in North America

Synopsis The latitudinal, regional, and annual variation in number of vertebrae and number of gill rakers present in sockeye salmon, Oncorhynchus nerka, stocks in North America was examined. Stocks in more northern areas had higher numbers of vertebrae and gill rakers than did those in more southern ones. Significant annual variability in the frequencies of these meristic characters within stocks was observed. When stocks were grouped into three regions (southern, central, and northern), heterogeneity in vertebral and gill raker frequencies was greater among regions than among stocks within the regions. Similarly, heterogeneity was greater among stocks than among sampling years within stocks. Differences in vertebral and gill raker frequencies are only useful for stock identification of sockeye salmon on a broad regional basis.     

Alewife,rainbow smelt and native fishes in Lake Michigan: competition or predation?

Synopsis Several native fish species in Lake Michigan became rare or locally extinct during the increase of rainbow smelt (1930s) and alewife (1950s–1960s). These particular native species have pelagic eggs or larvae which were large relative to the zooplankton and which co-occurred with feeding alewife, smelt or both. Alewife, smelt and most of the other planktivores in the lake probably consumed eggs, and at least alewife and smelt are known to consume larvae. For each species, I comment on the probable role of predation on early life history stages in their decline. Several species which currently co-exist with alewife and smelt have shown large scale declines during the increase of the exotics. Recruitment declines were often dramatic and the probability of predation on these species during their early life history is evaluated.     

Long-term zooplankton composition data reveal impacts of invasions on community composition in the Waikato lakes,New Zealand

Data collected on zooplankton community composition over longer time periods (>?10 years) are rare. We examined among-lake spatial and temporal trends of zooplankton communities from a monitoring programme undertaken in the Waikato region, New Zealand. A total of 39 lakes were sampled over a period of 12 years, between 2007 and 2019, with varying degrees of temporal effort. We focussed particularly on eight lakes, considered here as ‘long-term lakes’, where samples were collected with greater regularity (including 5 with 12 years of data). Among lakes, suspended sediment concentrations and indicators of lake trophic state were inferred to be important in determining the zooplankton distributions; as this region is dominated by shallow lakes, the relative importance of suspended sediments was high. Among the long-term lakes, the greatest dissimilarities in zooplankton community composition among years were in Lake Waahi, where the Australian Boeckella symmetrica was first detected in 2012. That is, the greatest temporal changes to zooplankton composition during the study period were due to the invasion by non-indigenous species, rather than changes in trophic state or other environmental variables; non-native species commonly dominated the individual counts of species through much of 2014 and 2015, with most samples since 2016 being again dominated by native species. Following this lake, the largest and shallowest lakes in the dataset—Whangape and Waikare—exhibited the greatest variability in community composition among years.

     

Genetic differentiation in gill raker number and length in sympatric anadromous and nonanadromous morphs of sockeye salmon, Oncorhynchus nerka

The genetic and environmental basis for polymorphism in gill raker number and length in sympatric anadromous and nonanadromous morphs of sockeye salmon, Oncorhynchus nerka, was investigated. Analysis of 30 full sib families involving pure types and reciprocal hybrids revealed that the variation was partitioned significantly among families within cross types and among cross types in both traits. As in the wild, kokanee displayed more gill rakers than sockeye; reciprocal hybrids displayed intermediate counts. Gill raker length also varied markedly among cross types, with pure sockeye displaying 19% longer gill rakers than comparable sized kokanee. This difference was in the opposite direction predicted, given the common positive association between gill raker number and length in sympatric morphs of the same species in fishes. Gill raker number and length were generally not correlated within cross types, suggesting independent divergence of the traits. The results are discussed in relation to genetic and trophic divergence of the morphs and to factors selecting for differentiation in the two gill raker traits.