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eliminate their vulnerability to enemies through adaptation, both behavioral and <br />morphological (Fryer 1986). How do fishes reduce vulnerability to enemies, and how <br />can species introductions can be so devastating? <br />It has been known for centuries that: "... major aspects of species' lifestyles are <br />moulded by the impact of natural enemies..." (Jefferies and Lawton 1984), and even <br />the morphology of a species can be changed by this interaction. One of the most <br />interesting fish examples is predator- induced morphological change in crucian carp <br />( Carassius carassius), whose populations increase their body depth in the presence of <br />northern pike to avoid being eaten (Bronmark and Minor 1992). This morphological <br />change is so dramatic that affected individuals have been mistakenly identified as a <br />separate species. <br />It takes time to develop the strategies that adapt one species to the activities of <br />others, and once developed, these strategies can be made less effective with physical or <br />biological alterations of habitat. Evolution usually occurs slowly (i.e., over geologic time <br />scales), but abrupt changes such as caused by human activities can disrupt the <br />relationships previously developed between species. For example, a reduction in <br />turbidity in the Missouri River could be expected to favor predation by native sauger on <br />chubs, sturgeon, or other species adapted to life in turbid water. However, the <br />introduction of nonnative fishes that may be "preadapted" to life in the altered <br />environments may place native predator and prey at a disadvantage. In this case, lack <br />of exposure to the nonnative species also could result in a naivete' to predation (e.g., <br />see Johnson et al. 1993), perhaps due to different foraging tactics, and etc. In this case, <br />defense strategies that evolved in response to sauger predation may not be appropriate <br />for introduced walleye, who are more adaptable, occupy different habitats, forage <br />differently, select different prey, and as a consequence are even known to displace <br />sauger in some habitats (Scott and Crossman 1973, Rawson and Scholl 1978). <br />The addition of other predators such as pike, bass, sunfishes, rainbow smelt, etc, to a <br />system that continues to be changed by physical habitat alteration makes it hard to <br />imagine how the native prey could survive for long. <br />But not all impacts caused by nonnative fishes are due to predation. Competition <br />for food and space also can be a powerful force (e.g., Schoenherr 1981, Taylor et al. <br />1984), especially with an increased vulnerability to predation. For example, a decrease <br />in turbidity can be expected to result in shifts in habitat use by predator -wary prey (Miner <br />and Stein 1996). In this case, less food and fewer places to hide greatly exacerbate <br />problems caused by increased predation and can result in species extirpations and <br />possibly extinctions over time. <br />A better understanding of vulnerability to other species can be aided by an <br />appreciation of interactions that develop by coevolution (i.e., evolution of 2 interacting <br />species that adapt to characteristics of the other and each enhances the fitness of the <br />other). In this case, coevolution can be considered "reciprocal evolution" among <br />22 <br />