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1948 <br />JOHNSON ET AL. <br />objectively rank-the threat posed by the three nonnative <br />predators based on relative consumptive demands. <br />Determining the predatory threat posed by a fish <br />population cannot be gauged simply by its abundance, <br />size structure, or characteristic dietary preferences. <br />Rather, we have shown how a number of factors <br />interact to determine a given species' capacity to harm <br />native fish populations; these factors include prey <br />choice, population size structure, abundance, and <br />physiological attributes and environmental suitability. <br />Bioenergetics models integrated all of these factors to <br />provide direct impact estimates in the form of the <br />consumptive demand for native fishes by each predator <br />population. These consumption estimates quantified <br />the current relative predatory threat posed by each <br />nonnative species, providing managers with hard <br />evidence with which to prioritize control efforts. <br />Despite their moderately high abundance, channel <br />catfish in the Yampa River contributed only about 1% <br />of total piscivory because their diet contained few fish. <br />Channel catfish were the most gape limited of the three <br />piscivores in our study; the gape width of a 400-mm <br />channel catfish was only about 77% of the gape width <br />of a 400-mm smallmouth bass (B.M.J., unpublished <br />data). Low piscivory is consistent with findings of <br />Tyus and Nikirk (1990), who reported that channel <br />catfish in the Yampa River rarely ate fish, even when <br />small-bodied fishes were abundant. In that study, only <br />large channel catfish (mean TL = 392 mm) were found <br />to have consumed any fish. Brooks et al. (2000) found <br />that piscivory by channel catfish was also low in the <br />San Juan River, Colorado-New Mexico-Utah, where <br />the prey assemblage was similar to that of the Yampa <br />River. Crayfish and insects were the predominant food <br />source of channel catfish in the Yampa River. <br />Competition for food between channel catfish and <br />native fishes has been put forward as a rationale for <br />channel catfish control efforts. Based on comparisons <br />of our channel catfish diet data with diet studies of <br />native fishes elsewhere (Vanicek and Kramer 1969; <br />Karp and Tyus 1990; Quist et al. 2006), considerable <br />diet overlap between channel catfish and native species <br />probably exists in the Yampa River. However, biomass <br />and production of invertebrate prey organisms in the <br />Yampa River are unknown; hence, resource limitation <br />and competition for food among native and nonnative <br />fishes cannot be inferred. <br />On aper-capita basis, northern pike consumed more <br />fish than the other predators in both the realized and <br />potential piscivory scenarios; this result is attributable <br />to the large body size of northern pike, their preference <br />for piscine prey, and the suitability of environmental <br />temperatures. However, because northern pike abun- <br />dance was relatively low, population-scale consump- <br />tion of fish by northern pike was similar to that by <br />smallmouth bass even though the smallmouth bass diet <br />contained a much smaller percentage of fish. Because <br />they were far less gape limited than smallmouth bass, <br />northern pike were able to maintain a relatively high <br />fraction of f sh in their diets after small-bodied fishes <br />declined; thus, northern pike were able to prey on <br />older, larger individuals that were still relatively <br />common in the system (Anderson 2005). Northern <br />pike are potent piscivores, capable of ingesting adults <br />of even large-bodied native species; therefore, continu- <br />ing efforts to reduce northern pike numbers in riverine <br />habitats where they prey on native species are <br />warranted. <br />We believe that the small contribution of fish to the <br />smallmouth bass diet and the incidence of cannibalism <br />in our data suggest that piscivory in the Yampa River <br />was limited by low availability of fish prey within the <br />gape limit of smallmouth bass. Spatial overlap between <br />small-bodied fishes and smallmouth bass (and northern <br />pike) is high all year long, and there are no <br />microhabitats that serve as refuges for native fishes, <br />Total piscivory by the smallmouth bass population was <br />similar to that by northern pike simply because <br />smallmouth bass were more abundant than northern <br />pike. smallmouth bass were highly piscivorous in the <br />GVR, where the invasion was more recent and where <br />small-bodied fishes were common (Bundy and Bestgen <br />2001). Based on potential piscivory scenarios, if <br />suitably sized fish prey were available in the Yampa <br />River, then smallmouth bass predation could-have been <br />10-fold higher than that of the other two predators <br />combined. Thus, the Yampa River smallmouth bass <br />population possessed a considerable level of latent <br />piscivory. Our analysis indicates that smallmouth bass <br />and northern pike each pose a serious threat to native <br />fishes but that smallmouth bass have the greatest <br />capacity to hamper native fish recovery in the Yampa <br />River by virtue of their high abundance. The <br />synergistic effects of the two predators may be <br />particularly devastating, because prey fish that manage <br />to outgrow the gape of smallmouth bass will remain <br />vulnerable to northern pike for years or a lifetime, <br />depending on the species. <br />Historic data on the fish assemblage in the Yampa <br />River before either northern pike or smallmouth bass <br />were abundant suggest that small-bodied fish density <br />was similar to present-day density in the GVR. Wick et <br />al. (1985) performed seining in shoreline and backwa- <br />ter habitats (n = 1,828 seine hauls) in the Yampa River <br />during 1981 and 1982. Mean density of mostly native, <br />small-bodied fishes for the 2 years combined was <br />34,000 fish/ha at RKM 165-191 and 45,000 fish/ha <br />over our entire study area (RKM 80-191). In the GVR, <br />