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7/14/2009 5:01:47 PM
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UCREFRP
UCREFRP Catalog Number
9511
Author
Portz, D. E. and H. M. Tyus
Title
Fish humps in two Colorado River fishes
USFW Year
2004
USFW - Doc Type
a morphological response to cyprinid predation?
Copyright Material
YES
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241 <br />were merely interested in drag and lift components <br />of fish body shapes with large nuchal humps. We <br />believe that the casts (rigid-body analogy) we used <br />were adequate for determining the role that pro- <br />truding rigid structures (nuchal humps) have on <br />drag and lift generated by fish that would be <br />attempting to hold their position in flowing water. <br />Thrust values were not sought after in this study; <br />rather, we focused on dead-drag. The use of dead- <br />drag is justified because we evaluated rigid struc- <br />tures and were not concerned with the action of <br />the fleshy body or turbulence caused by fins and <br />other structures (e.g., see Webb 1975). <br />Our use of rigid body casts instead of fresh <br />specimens may be criticized. However, the main <br />objective of this study was to evaluate whether a <br />hydrodynamic advantage might be induced solely <br />by large nuchal humps. Nuchal humps of G. cypha <br />and X. texanus are rigid body structures of bone <br />and hard muscle that have been thought to aid <br />these fishes hydrodynamically while attempting to <br />hold their position in a stream (Miller 1946, La <br />Rivers 1962). Thus, we used rigid casts to evaluate <br />the effect of humps on drag and lift in order to <br />avoid turbulence and higher drag coefficients that <br />would result as an artifact of body fluttering or due <br />to fins that may cause spurious readings in fresh <br />specimens (Webb 1975). Also, we did not address <br />shape-specific effects of humped fishes of different <br />sizes. No doubt, an evaluation of the allometric <br />relationships between nuchal hump size and the <br />drag and lift produced would provide useful <br />information about the functional consequences of <br />developing enlarged nuchal humps. However, <br />determining these relationships was beyond the <br />scope and budget of our study. This study is pri- <br />marily concerned with determining whether large <br />nuchal humps provide a hydrodynamic advantage <br />(i.e., negative lift with marginal drag), and offering <br />an alternate hypothesis of their usefulness. <br />The relatively high metabolic cost of swimming <br />is indisputable and a direct reflection of the drag <br />that must be overcome. For mobile organisms <br />such as stream fishes, the cost of excessive drag is <br />extreme, depending on habitats utilized. Fishes <br />living in swift water must expend energy at a rate <br />that is at least that of the product of drag and <br />velocity (Vogel 1994). Our analysis of drag on the <br />body casts of different fishes we studied produced <br />results that are in agreement with habitat prefer- <br />ences obtained by empirical studies: G. cypha and <br />X. texanus prefer slower habitats such as eddies, <br />backwaters and slackwaters (Tyus 1987, Karp & <br />Tyus 1990, Sigler & Sigler 1996). However, closely <br />related forms utilize high velocity habitats. In the <br />case of G. cypha, a congener, G. robusta, is more <br />streamlined, has a much less pronounced nuchal <br />hump, and does occur in swift currents. With re- <br />spect to X. texanus, its close relative, C. latipinnis, <br />also is more streamlined, lives in riffles and has no <br />hump. For the hydrodynamic advantage theory to <br />be correct, all other main channel fish species <br />should have a large hump to benefit swimming <br />performance in swift currents. Our findings show <br />that Colorado River fishes with the most stream- <br />lined bodies are fishes that historically inhabited <br />fast-flowing main river channels, whereas fishes <br />with large humps are known to occupy eddies and <br />slackwaters. In the case of X. texanus, a fish that <br />has been occasionally observed feeding in shallow <br />runs (Tyus 1987), enlarged pectoral fins may be an <br />adaptation to compensate for lift created by the <br />hump when the fish ventures into faster flows. <br />An apparently high energetic cost would be <br />associated with hump growth and locomotion with <br />the presence of a hump due to the drag that is <br />produced (Pettersson & Bronmark 1999). Al- <br />though it might be argued that humps are relict <br />structures that once had some purpose that has <br />been lost, logic dictates that some benefit must <br />accrue. One possible explanation, in view of our <br />findings that the humps do not aid swimming or <br />produce `negative' lift, might be sexual dimor- <br />phism as an aid in sexual selection. However, <br />males and females of both species produce humps; <br />thus, we discount sexual selection as the primary <br />factor in hump formation. Humps of males do <br />appear slightly larger, but we attribute such <br />dimorphism to building on an already existing <br />structure rather than developing the hump for this <br />purpose de novo. This is supported by a regression <br />of nuchal hump size vs. fish total lengths for fishes <br />that we measured at Dexter National Fish <br />Hatchery. In this case, no sexual dimorphism was <br />evident, and a smooth transition occurred as hump <br />size increased with fish growth (rz = 0.984 for X. <br />texanus; Figure 5 and r2 = 0.986 for G. cypha; <br />Figure 6). An additional possibility is that en- <br />larged nuchal humps are produced in these two <br />species in response to predation.
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