Intraspecific taxonomy and ecology characterize morphological divergence among cutthroat trout (Oncorhynchus clarkii ssp. Richardson) populations

We compared the proportion of morphological variation accounted for by subspecies categories with the proportion encompassed by ecologically based categories in cutthroat trout ( Oncorhynchus clarkii ssp.), as a means of assessing the relative importance of each approach in identifying intraspecific diversity. We used linear and geometric morphometrics to compare measures of body shape, fin length, and head features between and within subspecies of cutthroat trout. Both categories accounted for a significant proportion of the variation between and within the subspecies; however, the larger proportion was explained by subspecific differences, with the greatest morphological divergence between coastal cutthroat trout ( Oncorhynchus clarkii clarkii ) and interior subspecies. Ecotypic categories within each subspecies also explained significant morphological differences: stream populations had longer fins and deeper, more robust bodies than lake populations. The largest ecotypic differences occurred between stream and lake populations of Yellowstone cutthroat trout ( Oncorhynchus clarkii bouvieri ). Given that many cutthroat trout subspecies are of conservation concern, our study offers a better understanding of intraspecific variation existing within the species, providing precautionary evidence of incipient speciation, and a framework of describing phenotypic diversity that is correlated with ecological conditions.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 266–281.     

Morphological differentiation among local trout (Salmo trutta) populations

The trout (Salmo trutta) has been divided into three forms: sea-run trout, lake-run brown trout, and resident brown trout. They differ in their living environment, migratory behaviour, growth and appearance. As local trout populations are often isolated, and gene flow between them is minimal, differentiation between populations can be expected. The morphology of 1-year-old trout from ten populations representing all three forms was studied in a common-garden experiment. The fish were reared under similar environmental conditions, and 20 morphometric characters were measured from each individual fish. Marked morphological differentiation was found, and differences between populations were greater than differences between forms. The results suggest that the differences have a genetic basis, and they are likely to indicate adaptation to local environmental conditions in the native habitat of the trout.     

Effects of habitat and food resources on morphology and ontogenetic growth trajectories in perch

Studies on resource polymorphism have mainly been considered at the end stage of ontogeny, whereas many species undergo diet changes as they grow. We conducted a field survey to analyze the role of adaptive variation during ontogeny in Eurasian perch (Perca fluviatilis). We caught perch from the littoral and pelagic zones of a lake to investigate whether perch differ in morphology and diet between these habitats. We also investigated whether there were any differences in morphological trajectories during the ontogeny of perch collected from the two habitats. We found that perch caught in the littoral habitat, independently of size, had a deeper body, larger head and mouth and longer fins than perch caught in the pelagic zone. Macroinvertebrates and fish dominated the diet of littoral perch, whereas the diet of the pelagic perch consisted mainly of zooplankton and to some extent fish. Independently of size, the more streamlined individuals had a larger proportion of zooplankton and a smaller proportion of macroinvertebrates in their diet than the deeper-bodied individuals, indicating a relation between diet and morphology. Some morphological characters followed different ontogenetic trajectories in the two habitats; e.g. the changes to a deeper body and a larger head were faster in the littoral than in the pelagic perch. The relationship between the length of perch and the size of the mouth and fins also differed between perch from the two habitats, where the increase in the length of the pelvic fin and the area of the mouth increased faster with size in the littoral perch. Our findings show that variation in morphology between habitats differs during ontogeny in a way that corresponds to functional expectations for fish species that occupy these habitats.     

Morphological Consequences of Developmental Plasticity in <Emphasis Type="Italic">Rana temporaria</Emphasis> are not Accommodated into Among-Population or Among-Species Variation

Environmental induced developmental plasticity occurs in many organisms and it has been suggested to facilitate biological diversification. Here we use ranid frogs to examine whether morphological changes derived from adaptive developmental acceleration in response to pool drying within a species are mirrored by differences among populations and across species. Accelerated development in larval anurans under pool drying conditions is adaptive and often results in allometric changes in limb length and head shape. We examine the association between developmental rate and morphology within population, among populations in divergent environments, and among species inside the Ranidae frog family, combining experimental approaches with phylogenetic comparative analyses. We found that frogs reared under decreasing water conditions that simulated fast pool drying had a faster development rate compared to tadpoles reared on constant water conditions. This faster developmental rate resulted in different juvenile morphologies between the two pool drying conditions. The association between developmental rate and morphology found as a result of plasticity was not mirrored by differences among populations that differed in development, neither was it mirrored among species that differed in development rate. We conclude that morphological differences among populations and species were not driven by variation in developmental time per se. Instead, selective factors, presumably operating on locomotion and prey choice, seem to have had a stronger evolutionary effect on frog morphology than evolutionary divergences in developmental rate in the ranid populations and species studied.     

Juvenile current response, growth and maturity of above and below waterfall stocks of rainbow trout, Salmo gairdneri

Young rainbow trout were propagated from adults collected from above and below a water-fall, impassable to upstream fish migration on Kokanee Creek, tributary to Kootenay Lake, British Columbia. The two stocks of young (above and below waterfall) were reared in a hatchery under identical temperature, light and food conditions. Their directional response to water current was tested in a laboratory migration channel during their first summer and autumn. as well as the following spring. During tests in light, both stocks of young trout showed similar current responses, but in darkness the below falls stock showed much more downstream movement in the autumn than than did the above falls stock. In addition, below falls trout had a higher growth rate. were slightly heavier at the same length and matured earlier than the above falls stock. The adaptive significance of these differences are discussed in relation to maintenance of trout populations above waterfalls and to selective pressures faced by migratory below waterfall populations.     

Meristic variation among world hatchery stocks of rainbow trout,Salmo gairdneri Richardson

Synopsis Sixteen meristic characters were examined among 10 stocks of rainbow trout to establish their inter- and intrastock variability and relative discriminating power. The number of vertebrae, dorsal fin rays, scales above and below and lateral line, oblique scale rows and pectoral fin rays were found to be the most useful meristic characters in identifying stocks. Examination of the extent of meristic divergence among 10 hatchery or naturalized stocks from various regions of the world, using analysis of variance, the generalized distance function and per cent overlap, revealed that each stock exhibited highly significant differences in at least one meristic character, thereby demonstrating a high degree of intraspecific variability in meristic expression.In multivariate terms, stocks from Idaho and Alaska were almost totally divergent from all other stocks examined. Multivariate analysis of meristic features proves to be a useful method for identifying the extent of phenotypic variability among and within divergent stocks of rainbow trout. The observed meristic variability in test stocks of hatchery fish is seemingly genetically set within the limits previously described for native populations of rainbow trout.     

State of gonads of yearlings of triploid rainbow trout Oncorhynchus mykiss exposed to androgenic hormone under high-mountain conditions of South Vietnam

Changes in the morphology of gonads under the effect of methyltestosterone in juveniles of triploid rainbow trout Oncorhynchus mykiss incubated and cultivated in a trout farm under climatic conditions of South Vietnam have been studied. It was shown that the effect of the androgenic hormone intensifies the formation and development of testes in triploid rainbow trout at an age of 1+, decreasing the proportion of sterile individuals and the proportion of fish with anatomical and cytological anomalies in the structure of gonads. The relationship between the found anomalies in the structure of the genital system of trout and its triploidization, hormonal effect, and abiotic environmental conditions is discussed.     

Morphometric examination of the gills of walleye, Stizostedion vitreum vitreum (Mitchill) and rainbow trout, Salmo gairdneri Richardson

Observations on the gill morphology of individual gill arches of walleye, Stizostedion vitreum vitreum and rainbow trout, Salmo gairdneri suggest that the first two arches account for the highest proportion of gill filament number, secondary lamellae number, lamellae area, and gill surface area. Interspecific comparisons suggest walleye contain a larger number of gill filaments, with a lower secondary lamellae count, but a larger gill surface area than a trout of the same weight. This is partially attributed to the larger surface area of each lamella in a walleye than in a trout.
A detailed examination suggests the surface area of a lamella is dependent on its position on the gill filament, and the position of the gill filament on the gill arch.     

A COMPARISON OF TWO STICKLEBACKS

We present results of an experiment designed to address fundamental issues in the ecology and evolution of plastic trophic morphology: (1) Is observed plasticity adaptive? (2) How much interspecific morphological variation is the result of plasticity? (3) Have different selective regimes resulted in the evolution of different degrees of plasticity? (4) Is genetic variation for phenotypic plasticity present in contemporary populations? We raised fish from two recently diverged species of freshwater threespine sticklebacks on two different diets representative of the natural prey of the two species. Both species exhibited morphological plasticity in an adaptive direction: each species more closely resembled the other when raised on the latter's diet. Dietreversal reduced the natural morphological gap between these two species, -1% to 58%, depending on the trait. One species is known to have a more variable diet in the wild than the other species, and we found that it also exhibited the greater amount of morphological plasticity. Given that the two species have recently diverged, this result is compelling evidence that diet variability is important in the evolution of plastic trophic morphology. Finally, by using a full-sib experimental design, we demonstrated that genetic variation for morphological plasticity exists in contemporary populations, thus confirming that plasticity has evolutionary potential.     

Habitat use by Nonnative Rainbow Trout, Oncorhynchus mykiss, and Native Little Colorado spinedace, Lepidomeda vittata

We evaluated overlap in microhabitat use between nonnative rainbow trout, Oncorhynchus mykiss, and native Little Colorado spinedace, Lepidomeda vittata, a federally threatened cyprinid, in natural and experimental settings. In natural settings, we also examined occurrence and microhabitat use of two other native fishes, speckled dace, Rhinichthys osculus, and bluehead sucker, Catostomus discobolus. Native species co-occurred, as did rainbow trout and bluehead sucker. However, occurrences of Little Colorado spinedace and speckled dace were not significantly correlated with occurrence of rainbow trout. Total lengths of all three native species were significantly smaller at allopatric sites than at sites sympatric with rainbow trout. Microhabitat characteristics at sites with rainbow trout did not differ from those where the other three species were found, but did differ among the native species. In laboratory experiments with Little Colorado spinedace and rainbow trout, rainbow trout used the lower depth zone most, and spinedace increased use of the lower depth zone upon addition of rainbow trout. In addition, species tended to co-occur in zones, but used cover independently of one-another, suggesting a low level of agonistic interactions. However, after addition of a high density of rainbow trout, spinedace tended to use cover less than before. We suggest that the species can coexist at low rainbow trout densities. Potential negative effects of rainbow trout on Little Colorado spinedace likely increase with increasing densities of rainbow trout, and rainbow trout likely affect smaller size classes of Little Colorado spinedace more than larger ones.     

Differences in Seed Production Between Stellaria media Populations from Different Habitat Types

Seed production in several populations of Stellaria media fromdifferent types of habitat was examined. Two populations, onefrom a flower garden and the other from a herring gull colonywere compared in detail by growing them at three soil fertilitylevels under similar conditions in a greenhouse. Three important differences between the two populations independentof soil fertility emerged: the garden plants flowered on averageeleven weeks earlier, they produced much smaller seeds, andthey produced more seeds per capsule. Other differences were influenced by soil fertility: at highfertility the garden plants produced about 50 per cent moreseeds than the gull colony plants, but on a seed weight basis,the gull colony plants were the more productive. However forboth populations at high fertility the proportion of shoot biomassallocated to seed production was similar. Some of these differences can be accounted for in terms of conditionsoccurring in the two habitats, in particular the relative importanceof disturbance and competition at the sites. Chickweed, Stellaria media (L.) Vill., ecotypic differentiation, time to flowering, seed production, resource allocation     

Intraspecific Trait Variation Driven by Plasticity and Ontogeny in Hypochaeris radicata

The importance of intraspecific variation in plant functional traits for structuring communities and driving ecosystem processes is increasingly recognized, but mechanisms governing this variation are less studied. Variation could be due to adaptation to local conditions, plasticity in observed traits, or ontogeny. We investigated 1) whether abiotic stress caused individuals, maternal lines, and populations to exhibit trait convergence, 2) whether trait variation was primarily due to ecotypic differences or trait plasticity, and 3) whether traits varied with ontogeny. We sampled three populations of Hypochaeris radicata that differed significantly in rosette diameter and specific leaf area (SLA). We grew nine maternal lines from each population (27 lines total) under three greenhouse conditions: ambient conditions (control), 50% drought, or 80% shade. Plant diameter and relative chlorophyll content were measured throughout the experiment, and leaf shape, root∶shoot ratio, and SLA were measured after five weeks. We used hierarchical mixed-models and variance component analysis to quantify differences in treatment effects and the contributions of population of origin and maternal line to observed variation. Observed variation in plant traits was driven primarily by plasticity. Shade significantly influenced all measured traits. Plant diameter was the only trait that had a sizable proportion of trait variation (30%) explained by population of origin. There were significant ontogenetic differences for both plant diameter and relative chlorophyll content. When subjected to abiotic stress in the form of light or water limitation, Hypochaeris radicata exhibited significant trait variability. This variation was due primarily to trait plasticity, rather than to adaptation to local conditions, and also differed with ontogeny.     

Mitochondrial and nuclear DNA sequence variability among populations of rainbow trout (Oncorhynchus mykiss)

Mitochondrial and nuclear DNA variability was examined to assess population genetic structure and phylogeographic relationships in rainbow trout. Single-strand conformation polymorphisms and restriction site differences within 1055 bp of the mitochondrial D-loop region and 1566 bp of nuclear DNA in six single-copy nuclear DNA regions identified 31 mitochondrial genotypes and 50 nuclear alleles. Gene trees were constructed by sequencing each variant allele or mitochondrial genotype identified. Examination of 30 populations in 10 native rainbow trout groups using an analysis of molecular variance ( AMOVA ) indicated that 65% of mitochondrial variability and 35% of nuclear variability was explained by differences among the 10 groups. Phylogenetic patterns evident in mitochondrial and nuclear DNA were not always concordant. Differences in the evolutionary patterns detected by mitochondrial and nuclear DNA may reflect the differential impact of past introgression events on variability in the two genomes.     

Factors influencing return rate and marine residence duration in sea trout populations in Central Norway

Brown trout (Salmo trutta) display extensive plasticity in marine migratory behaviours, with marine migrations considered to be an adaptive strategy which enables sea trout to maximize growth and reproductive potential. However, marine migrations are not without associated costs, including threats posed by ever-increasing salmon lice (Lepeophtheirus salmonis) infestations. In the present study, we used passive integrated transponder technology to characterize variability in sea trout migration behaviour amongst three catchments situated in a region of intensive salmon farming in central Norway. Specifically, we investigate how lice infestation, out-migration date and body size alter sea trout return rate and marine residence duration during the first out-migration to sea from each catchment. Distinct catchment-specific differences in sea trout out-migration size and the number of cohorts were observed, but larger body size did not guarantee the successful return of migrating trout. The marine residence duration of individuals that successfully returned to freshwater was positively correlated with lice infestation risk, suggesting for these individuals the lethal infestation threshold had not been reached. Our results also suggest that sea trout populations from lotic-dominated catchments are potentially at greater risk from size-related threats to their survival encountered during their marine migrations than sea trout from lentic-dominated catchments. The variability in sea trout migratory behaviour amongst catchments observed here emphasizes the challenges fisheries managers face when deciding the best actions to take to protect the anadromous portion of brown trout populations.     

Ecotype differentiation and coexistence of two parapatric tetraploid subspecies of cocksfoot (Dactylis glomerata) in the Alps

Two tetraploid subspecies of Dactylis glomerata L., subsp. reichenbachii (Hausm.) Stebbins et Zohary and subsp. glomerata , occur in the French Alps. The former is confined to dolomitic, south-facing, alpine lawns above 2000 m, whereas the latter occurs in non-dolomitic habitats in subalpine meadows mainly below 1900 m. Previous studies of allozyme variation have shown that genetic introgression between the two subspecies occurs over large areas. By contrast, morphologically intermediate individuals only occur in an extremely narrow area, suggesting that the morphological and physiological differences between the two subspecies is of adaptive significance. A reciprocal clone transplant experiment was set up to examine (1) any genetic differences between subspecies indicative of ecotypic differentiation in relation to habitat characteristics and (2) the level of phenotypic plasticity in the two subspecies. Genetic differentiation was confirmed by a statistically significant taxon × site interaction effect in anova for all traits studied. The glomerata populations produced more tillers, longer leaves and higher culms in all sites, especially in their home environment. However, reichenbachii populations produced more seeds than the glomerata populations in the original reichenbachii environment, suggesting ecotypic differentiation between the two subspecies. This result might also explain why the glomerata subspecies is unable to colonize dolomitic habitats occupied by the reichenbachii subspecies. The reichenbachii populations showed less plasticity than the glomerata populations for leaf length and floriferous tiller number, a result which is discussed in the context of the response of plants from productive and non-productive habitats to environmental variation.     

Phenotypic plasticity inPotamogeton (Potamogetonaceae)

Sources of the extensive morphological variation of the species and hybrids ofPotamogeton were studied, especially from the viewpoint of the stability of the morphological characters used inPotamogeton taxonomy. Transplant experiments, the cultivation of clones under different values of environmental factors, and the cultivation of different clones under uniform conditions were performed to assess the proportion of phenotypic plasticity in the total morphological variation. Samples from 184 populations of 41Potamogeton taxa were grown. The immense range of phenotypic plasticity, which is possible for a single clone, is documented in detail in 14 well-described examples. The differences among distinct populations of a single species observed in the field were mostly not maintained when grown together under the same environmental conditions. Clonal material cultivated under different values of environmental factors produced distinct phenotypes, and in a few cases a single genotype was able to demonstrate almost the entire range of morphological variation in an observed trait known for that species. Several characters by recent literature claimed to be suitable for distinguishing varieties or even species were proven to be dependent on environmental conditions and to be highly unreliable markers for the delimiation of taxa. The unsatisfactory taxonomy that results when such classification of phenotypes is adopted is illustrated by three examples from recent literature. Phenotypic plasticity was found to be the main source of morphological variation within the species ofPotamogeton, having much more influence than morphological differences caused by different genotypes.     

Which Extent is Plasticity to Light Involved in the Ecotypic Differentiation of a Tree Species from Savanna and Forest?

Light intensity and heterogeneity are some of the main environmental factors that differ between forest and savanna habitats, and plant species from these habitats form distinct functional types. In this study, we tested the hypothesis that not only differences in morphological and physiological traits but also phenotypic plasticity in response to light are involved in adaptation to forest and savanna habitats by investigating ecotypic differentiation between populations of Plathymenia reticulata (Leguminosae: Mimosoideae), a tree from the Brazilian Atlantic Forest and the Brazilian Cerrado (savanna). Seeds from four natural populations (one from each biome core area and two from ecotonal regions) were grown in a common garden with four light treatments. Fifteen morphological and physiological characteristics were evaluated until individuals reached 6 mo old. Comparisons among populations showed differences for seven traits in at least one light treatment. These differences pointed to local adaptation to different biomes. Populations showed different levels of phenotypic plasticity in response to light in seven traits. Higher plasticity was found either in the forest core population or ecotonal populations; lower values were found in the cerrado core population. Lower plasticity in the cerrado population emphasizes the stress resistant syndrome, as lower plasticity is probably advantageous in a habitat where a conservative resource use is crucial. Higher plasticity in forest individuals suggests higher ability in exploiting the light heterogeneity in this habitat. Also, higher plasticity in ecotonal populations can be important to ensure the maintenance of P. reticulata in these temporally and spatially dynamic areas. Abstract in Portugese is available at http://www.blackwell‐synergy.com/loi/btp .     

Is genetic variability so important? Non-native salmonids in South America

Three salmonid species introduced in Patagonian national parks in Argentine have experienced different degrees of expansion. Atlantic salmon Salmo salar is restricted to a few river-lake systems and its populations have been declining over recent years. Both rainbow Oncorhynchus mykiss and brown trout Salmo trutta populations have expanded from their introduction sites and now occupy a wide range of freshwater ecosystems. Genetic variation at the same neutral markers (microsatellite loci) was examined for different populations of the three species acclimatized to the same areas, and compared with that of native populations. Founder effects denoted as reduced variability and great differentiation with respect to the native populations were detected. Significant reduction in variability has not been an obstacle for successful adaptation of rainbow and brown trout, indicating that genetic variability per se cannot be claimed as the reason for their different outcomes in the new habitats.     

Effects of species invasion on population dynamics,vital rates and life histories of the native species

Invasions occurring in natural environments provide the opportunity to study how vital rates change and life histories evolve in the presence of a competing species. In this work, we estimate differences in reproductive traits, individual growth trajectories, survival, life histories and population dynamics between a native species living in allopatry and in sympatry with an invasive species of the same taxonomic Family. We used as a model system marble trout Salmo marmoratus (native species) and rainbow trout Oncorhynchus mykiss (non-native) living in the Idrijca River (Slovenia). An impassable waterfall separates the stream into two sectors only a few 100 meters apart: a downstream sector in which marble trout live in sympatry with rainbow trout and an upstream sector in which marble trout live in allopatry. We used an overarching modelling approach that uses tag-recapture and genetic data (>2,500 unique marble and rainbow trout were sampled and genotyped) to reconstruct pedigrees, test for synchrony of population dynamics and model survival and growth, while accounting for individual heterogeneity. The population dynamics of the two marble trout populations and of rainbow trout were synchronous. We found higher prevalence of younger parents, higher mortality and lower population density in marble trout living in sympatry with rainbow trout than in marble trout living in allopatry. There were no differences in the average individual growth trajectories between the two marble trout populations. Faster life histories of marble trout living in sympatry with rainbow trout are consistent with predictions of life history theory.     

Hybridization dynamics between Colorado's native cutthroat trout and introduced rainbow trout

Newly formed hybrid populations provide an opportunity to examine the initial consequences of secondary contact between species and identify genetic patterns that may be important early in the evolution of hybrid inviability. Widespread introductions of rainbow trout (Oncorhynchus mykiss) into watersheds with native cutthroat trout (Oncorhynchus clarkii) have resulted in hybridization. These introductions have contributed to the decline of native cutthroat trout populations. Here, we examine the pattern of hybridization between introduced rainbow trout and 2 populations of cutthroat trout native to Colorado. For this study, we utilized 7 diagnostic, codominant nuclear markers and a diagnostic mitochondrial marker to investigate hybridization in a population of greenback cutthroat trout (Oncorhynchus clarkii stomias) and a population of Colorado River cutthroat trout (Oncorhynchus clarkii pleuriticus). We infer that cutthroat-rainbow trout hybrid swarms have formed in both populations. Although a mixture of hybrid genotypes was present, not all genotype combinations were detected at expected frequencies. We found evidence that mitochondrial DNA introgression in hybrids is asymmetric and more likely from rainbow trout than from cutthroat trout. A difference in spawning time of the 2 species or differences in the fitness between the reciprocal crosses may explain the asymmetry. Additionally, the presence of intraspecific cytonuclear associations found in both populations is concordant with current hypotheses regarding coevolution of mitochondrial and nuclear genomes.