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235 <br />rate of removal of momentum from a flowing <br />fluid; Webb 1975, Vogel 1994). <br />Reynolds numbers indicate the kinds of forces <br />that have to be overcome by an object moving in a <br />fluid. These numbers are a ratio of the inertial and <br />viscous forces, which for fish are dependant on <br />three factors: the swimming velocity of the fish, <br />characteristic length of the fish (where shape is <br />factored in), and viscosity of the water (Webb <br />1975, Denny 1988). Streams exhibit Reynolds <br />number values of approximately 100 000, where <br />neither viscous nor inertial forces dominate (Webb <br />1975, Vogel 1994, Lampert & Sommer 1997), and <br />shapes of fishes that result in a hydrodynamic <br />advantage may sometimes be difficult to predict. <br />In most cases, stream fishes reduce structures that <br />induce drag because increased drag also would <br />increase energetic costs. Adult stream fishes are <br />generally streamlined, with an exception in fishes <br />that do not carry out sustained swimming (Webb <br />1984, Diana 1995). <br />In addition to drag, another hydrodynamic <br />consideration for life in moving fluids is lift. Lift <br />can be defined as any force that is normal to the <br />direction of flow, in this case up (i.e., `positive lift') <br />and down (i.e., `negative lift'; Denny 1988). The <br />amount of lift generated by morphology depends <br />on how much the flow is turned by the shape of the <br />object. If flow above and below the shape are the <br />same, no net lift results. However, if water pres- <br />sure on the dorsal surface is less than that below, <br />the result is a net upward force: in this case, the <br />increased area of the leading edge deflects the <br />water so that the streamlines are crowded together <br />above the object. This crowding thus causes an <br />increase in water velocity on the dorsal surface <br />resulting in dynamic lift (Schlichting 1979, Denny <br />1988, Vogel 1994). <br />The `nuchal hump hydrodynamic advantage <br />hypothesis' attributed to Miller (1946) and La <br />Rivers (1962) can be evaluated by determining the <br />components of drag and lift produced by the <br />humps. In this study we determine drag and lift <br />components associated with large humps to eval- <br />uate whether the humps confer a benefit for life in <br />fast flow. In addition, we explore an alternate <br />hypothesis regarding the advantage of large humps <br />by examining other factors exerting evolutionary <br />pressures for hump development, such as a need <br />for a morphological defense against predators. <br />Materials and methods <br />Drag and lift <br />Three of the fishes we studied, G. cypha, G. elegans <br />and X. texanus, are endangered species protected <br />by federal and state statutes, thus live or freshly <br />sacrificed fishes could not be used in hydrody- <br />namic tests. Instead, polyester resin casts of mu- <br />seum specimens were used to obtain information <br />on how different body forms would be affected by <br />water flow. <br />We made whole-body casts of each species from <br />silicone rubber molds. We suspended fish speci- <br />mens in a plexiglas molding box filled with GI- <br />1000 RTV Silicone Rubber (Silicones, Inc.). We <br />mixed the silicone rubber with a fixed amount of <br />activator, placed it into a vacuum chamber, and <br />rapidly degassed it before pouring it into the <br />molding box. Once rubberized, we removed the <br />mold from the Plexiglas® box, divided it into two <br />halves, and retrieved the fish from the mold. <br />The cast-making procedure was as follows: the <br />silicone rubber mold was placed back into the <br />Plexiglaso box to maintain shape and we placed a <br />brass mounting apparatus (used to connect the <br />two-dimensional force beam to the fish cast) inside <br />the mold. We then filled the mold with polyester <br />resin mixed with 60 pm diameter glass micro- <br />spheres (3M S60; Minnesota Mining and Manu- <br />facturing Company) to 40% of the total volume to <br />add strength to the casts and to serve as filler. <br />After a day of hardening time, we removed the <br />casts with embedded mounting apparatuses from <br />the molds and sanded them to remove any <br />imperfections. <br />We made casts from molds of different fish <br />morphotypes to evaluate the hydrodynamics of <br />each body plan. We made two different casts of <br />each species morphotype using a cast most repre- <br />sentative of a fully developed, mature adult. We <br />used casts from the fish exhibiting enlarged nuchal <br />humps (i.e., a 381 mm TL G. cypha and a 458 mm <br />TL X. texanus) as the treatment group to measure <br />drag and the direction of force resulting from the <br />prominent nape. We later removed the humps on <br />these fish models and retested the casts to assess <br />performance without the humps. We compared lift <br />and drag values for those casts with nuchal humps <br />to the same casts with humps removed. We used