Ernest R. Keeley

The origins of ecotypic variation of rainbow trout: a test of environmental vs. genetically based differences in morphology

Although morphological plasticity has been observed in a variety of taxa, few experimental studies have compared the relative proportion of morphological variability that is accounted for by environmentally induced plasticity, and how much is because of genetically based differences among populations. We compared the morphology of six rainbow trout (Oncorhynchus mykiss) populations from different ecotypic categories that were raised under flowing vs. standing‐water conditions. Our data indicate that both environmental conditions and ecotypic differences account for a significant proportion of variation in morphology. Among ecotype effects, however, accounted for a much larger proportion of morphological variability than environmental conditions. Rainbow trout from stream populations had deeper caudal peduncles, and longer fins than lake populations, and rainbow trout from a piscivorous population had larger mouth and head lengths than all other ecotypes. Environmentally induced differences in morphology were primarily related to differences in mouth and head lengths, as well as fin length. Relative to morphometric differences from natural rainbow trout populations, most characteristics deviated in the same direction in our experimental populations. Our data indicate that morphological differences across rainbow trout populations have a genetic basis and may represent locally adaptive characteristics and highlight the role of ecology in promoting phenotypic divergence.

Geographic patterns of introgressive hybridization between native Yellowstone cutthroat trout ( Oncorhynchus clarkii bouvieri) and introduced rainbow trout ( O. mykiss ) in the South Fork of the Snake River watershed, Idaho

Throughout its native range, the Yellowstone cutthroat trout (YCT), Oncorhynchus clarkii bouvieri, is declining dramatically in both abundance and distribution as a result of introgression with introduced rainbow trout (RBT), O. mykiss. We sampled over 1,200 trout from the South Fork of the Snake River (SFSR) watershed, in southeastern Idaho and western Wyoming, and measured the extent of introgression of RBT genes into native gene pools of YCT using seven species-specific, co-dominant nuclear genetic markers. We also used mitochondrial DNA (mtDNA) haplotype differences between the two parental trout species to determine the directionality of the hybridization. We found low levels of RBT introgression (only 7% of sampled individuals had one or more RBT alleles) into YCT gene pools, with the majority of hybrids (78%) occurring in mainstem localities of the SFSR and in lower elevation reaches of certain tributaries. Hybridization was bidirectional with respect to mtDNA haplotype, but the majority of hybrids (75%) had YCT maternal haplotypes, indicative of the greater proportion (90%) of YCT-genotypes in the SFSR watershed. The primary factor influencing the geographic distribution of RBT introgressed individuals was fluvial distance from localities of stocking origin. To a lesser extent, elevation, also influenced the distribution of hybrid genotypes, with several entire tributaries and all upper elevation reaches within tributaries harboring only YCT-genotypes. Important management implications of the study suggest targeting particular tributaries and upper reaches within tributaries for YCT protection and exclusion of RBT hybrid colonization.

Competition between native and introduced salmonid fishes: cutthroat trout have lower growth rate in the presence of cutthroat-rainbow trout hybrids.

When nonnative species become established within new communities, competition may play a role in determin- ing the persistence of ecologically similar native species. In western North America, many native cutthroat trout (Onco- rhynchus clarkii) populations have been replaced by nonnative rainbow trout (Oncorhynchus mykiss). Superior competitive ability of rainbow trout and cutthroat-rainbow trout hybrids is often cited for this replacement; however, few studies have tested for mechanisms that might allow introduced rainbow trout to out-compete native trout species. Our previous work found individual-based differences in swimming and foraging ability among cutthroat trout, rainbow trout, and their hy- brids. In this study, we tested for the presence and strength of competition between cohorts of cutthroat trout, rainbow trout, and their reciprocal hybrids. We assayed the growth rate of juvenile cutthroat trout in allopatry versus cutthroat trout when sympatric with rainbow trout and each hybrid cross. After controlling for size and density of trout, cutthroat trout co- horts in stream channels that contained hybrid genotypes experienced lower growth than cutthroat trout in allopatry. Aver- aged across heterospecific treatments, cutthroat trout growth was also lower than that of cutthroat trout cohorts in allopatry. Our study suggests that juvenile cutthroat trout experience a growth disadvantage when competing against cutthroat-rainbow hybrids.

Conservation prioritization in widespread species: the use of genetic and morphological data to assess population distinctiveness in rainbow trout (Oncorhynchus mykiss) from British Columbia, Canada

Prioritization of efforts to maintain biodiversity is an important component of conservation, but is more often applied to ecosystems or species than within species. We assessed distinctiveness among 27 populations of rainbow trout (Salmonidae: Oncorhynchus mykiss) from British Columbia, Canada, using microsatellite DNA variation (representing historical or contemporary demography) and morphology (representing adaptive variation). Standardized genetic scores, that is, the average deviation across individuals within populations from the overall genetic score generated by factorial correspondence analysis, ranged from 1.05 to 4.90 among populations. Similar standardized morphological scores, generated by principal components analysis, ranged from 1.19 to 5.35. There was little correlation between genetic and morphological distinctiveness across populations, although one population was genetically and morphologically the most distinctive. There was, however, a significant correlation (r = 0.26, P = 0.008) between microsatellite (FST) and morphological (PST) divergence. We combined measures of allelic richness, genetic variation within, and divergence among, populations and morphological variation to provide a conservation ranking of populations. Our approach can be combined with other measures of biodiversity value (habitat, rarity, human uses, threat status) to rationalize the prioritization of populations, especially for widespread species where geographic isolation across distinct environments promotes intraspecific variability.

Bioenergetic assessment of habitat quality for stream-dwelling cutthroat trout (Oncorhynchus clarkii bouvieri) with implications for climate change and nutrient supplementation

We used a bioenergetic model to determine if cutthroat trout (Oncorhynchus clarkii bouvieri) abundance was related to net energy intake rates (NEI) and the proportion of suitable habitat and to evaluate potential changes in habitat quality due to climate change and stream fertilization efforts. We conducted monthly sampling of cutthroat trout, invertebrate drift, and physical habitat features in pool and riffle habitats. Fish in this study selected foraging positions that enabled them to maximize NEI, and most fish were capable of sustaining high growth rates from July to September. Mean NEI and the proportion of suitable habitat at sites were greater in pools relative to riffle habitats and declined from July to October, primarily due to a decline in temperature over the four months. Cutthroat trout biomass was significantly related to NEI and the proportion of suitable habitat at a site. Model simulations indicated that climate change might reduce habitat quality for small-bodied trout, while extending ...

A comparison of aggressive and foraging behaviour between juvenile cutthroat trout, rainbow trout and F1 hybrids.

‘Successful’ introduced species are often thought to cause declines or extinctions of native species through competitive superiority. In western North America, introduced rainbow trout, Oncorhynchus mykiss, have completely replaced many native cutthroat trout, Oncorhynchus clarkii, populations; however, few studies have identified the mechanisms that may allow rainbow trout to outcompete cutthroat trout. We raised Yellowstone cutthroat trout, Oncorhynchus clarkii bouvieri, rainbow trout, and their first generation hybrids in a common environment and conducted pairwise contests to test for differences in aggression, ability to defend a feeding station, and amount of food captured between these species and their hybrids. We did not detect a difference in number of aggressive acts conducted between cutthroat, rainbow and hybrid trout; however, cutthroat trout had the lowest success in occupying the feeding station and captured a lower proportion of food than rainbow and hybrid trout. Furthermore, hybrid crosses and rainbow trout had highest success at holding the feeding station and capturing food items when competing against cutthroat trout. Our study suggests that juvenile Yellowstone cutthroat trout are less successful at maintaining profitable feeding territories and capturing food items when competing against rainbow trout and first generation hybrids.

Watershed boundaries and geographic isolation: patterns of diversification in cutthroat trout from western North America

BackgroundFor wide-ranging species, intraspecific variation can occur as a result of reproductive isolation from local adaptive differences or from physical barriers to movement. Cutthroat trout (Oncorhynchus clarkii), a widely distributed fish species from North America, has been divided into numerous putative subspecies largely based on its isolation in different watersheds. In this study, we examined mtDNA sequence variation of cutthroat trout to determine the major phylogenetic lineages of this polytypic species. We use these data as a means of testing whether geographic isolation by watershed boundaries can be a primary factor organizing intraspecific diversification.ResultsWe collected cutthroat trout from locations spanning almost the entire geographic range of this species and included samples from all major subspecies of cutthroat trout. Based on our analyses, we reveal eight major lineages of cutthroat trout, six of which correspond to subspecific taxonomy commonly used to describe intraspecific variation in this species. The Bonneville cutthroat trout (O. c. utah) and Yellowstone cutthroat trout (O. c. bouvieri) did not form separate monophyletic lineages, but instead formed an intermixed clade. We also document the geographic distribution of a Great Basin lineage of cutthroat trout; a group typically defined as Bonneville cutthroat trout, but it appears more closely related to the Colorado River lineage of cutthroat trout.ConclusionOur study indicates that watershed boundaries can be an organizing factor isolating genetic diversity in fishes; however, historical connections between watersheds can also influence the template of isolation. Widely distributed species, like cutthroat trout, offer an opportunity to assess where historic watershed connections may have existed, and help explain the current distribution of biological diversity across a landscape.

Temperature-Related Changes in Habitat Quality and Use by Bonneville Cutthroat Trout in Regulated and Unregulated River Segments

Abstract The availability of suitable habitat is often viewed as one of most important limiting factors for animal populations. In this study, we examined the composition and spatial distribution of stream habitat based on summer water temperature using airborne thermal imagery, floating temperature surveys, and fixed temperature data loggers in a regulated and unregulated segment of the Bear River in Idaho and Wyoming. We also used temperature-sensitive radio telemetry tags to measure water temperature in habitats used by Bonneville cutthroat trout Oncorhynchus clarkii utah. We found that when available water temperatures increased in the Bear River, cutthroat trout in the regulated segment continuously selected cooler habitats that followed a similar rate of increase as available water temperatures. In the unregulated segment, cutthroat trout did not select significantly cooler available water temperatures when measured over the entire study period. However, cutthroat trout in the unregulated segment di...

Home Range Size and Foraging Ecology of Bull Trout and Westslope Cutthroat Trout in the Upper Salmon River Basin, Idaho

Abstract We used radiotelemetry to determine how body size, foraging ecology, and seasonal activity influence the home range size of adult bull trout Salvelinus confluentus and westslope cutthroat trout Oncorhynchus clarkii lewisi in the upper Salmon River basin of central Idaho. Stomach samples were collected to determine diet composition and compare foraging strategies between the two species. Contrary to other studies of home ranges for stream-dwelling fishes, home range size was not significantly related to total length or body mass in either species. Although overall home range size did not differ between species, monthly movement (km) was significantly greater for bull trout than for westslope cutthroat trout during July and September. Stomach content analysis revealed that the size of prey consumed was significantly larger for bull trout than for westslope cutthroat trout but was not significantly correlated with body size in either species. Home range size for bull trout and westslope cutthroat tr...

Prey size, prey abundance, and temperature as correlates of growth in stream populations of cutthroat trout

Growth and maximum size of stream fishes can be highly variable across populations. For salmonid fishes in streams, individuals from populations confined to headwater streams often exhibit small size at maturity in comparison to populations with access to main-stem rivers. Differences in prey size, prey availability, and metabolic constraints based on temperature may explain patterns of maximum size and growth. In this study, cutthroat trout from headwater stream populations that were isolated above a waterfall were compared to individuals from populations in similar sized streams without a movement barrier and from large main-stem rivers. Cutthroat trout from smaller streams with or without a movement barrier were significantly smaller at a given age than fish from main-stem rivers, where individuals were able to achieve a much larger maximum size. Comparisons of invertebrate drift abundance and size in the three types of streams revealed that drift size did not differ between stream categories, but was highest per volume of water in large main-stem rivers. Across all stream types, prey abundance declined from summer to fall. Temperature declined over the course of the season in a similar manner across all stream types, but remained relatively high later in the season in main-stem river habitats. Prey availability and temperature conditions in main-stem rivers may provide more optimal growing conditions for fish as individuals increase in size and become constrained by prey availability and temperature conditions in small streams. Maintaining connectivity between small spawning and rearing tributary streams and main-stem river habitats may be critical in maintaining large-bodied populations of stream salmonids.