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234 <br />900 mm are exceedingly rare. The largest P. lucius <br />captured in the multiagency Upper Colorado <br />River Fishes Recovery Implementation Program <br />was recorded at 1240 mm TL and only 7 of the <br />more than 5000 P. lucius captured since the 1970s <br />have exceeded 900 mm TL (C. McAda, US Fish & <br />Wildlife Service, personal communication). The <br />historical record is too scant to make valid state- <br />ments about past fish sizes other than to speculate <br />about maximum sizes that were probably attained <br />(e.g., Miller 1961). Two other fish species of the <br />Colorado River, the razorback sucker, Xyrauchen <br />texanus, and the humpback chub, Gila cypher, have <br />been described as `bizarre' because they exhibit <br />large nuchal processes that are located dorsally, <br />posterior to the head. These pronounced humps <br />look similar but are not homologous structures. <br />The X. texanus hump is bony, formed mostly by <br />enlarged neural and interneural bones (La Rivers <br />1962), while the G. cypha hump consists of a <br />muscular mass (Miller 1946, Figure 1). <br />The function of these nuchal humps is un- <br />known, though Miller (1946) speculated that the <br />G. cypha hump provides a hydrodynamic advan- <br />tage in swift currents. Miller reasoned that water <br />flowing over the hump would produce a down- <br />ward force on the fish, aiding the fish in main- <br />taining position on the bottom. Flow at the <br />benthic boundary layer is slower and would re- <br />quire less exertion to swim or maintain position in <br />a river. Unfortunately, Miller (1946) was unable to <br />observe G. cypha swimming in nature and little <br />was known about its habitat use. La Rivers (1962) <br />offered a similar explanation for X. texanus, <br />hypothesizing that flowing water would push the <br />sloping head downward while the bony keel pro- <br />vided increased stability in fast flows, thus <br />steadying the fish against the bottom. However, La <br />Rivers (1962) also was unable to test his hypoth- <br />esis. Yet, their assumptions that nuchal humps are <br />hydrodynamically advantageous adaptations to <br />swift currents are often cited (e.g., Cole 1994, Si- <br />gler & Sigler 1996, Moyle & Cech 2004). <br />Additional information about the adaptations <br />and life histories of Colorado River fishes gathered <br />in the last 20 years has prompted us to reevaluate <br />the function of large nuchal humps. It is now <br />understood that humps in both species are formed <br />later in life at a time when adult G. cypha pre- <br />dominantly occupy eddies and adult X. texanus <br /> <br />C ? 6, <br />tl <br />Figure 1. Four endangered Colorado River fishes: (a) Xyrau- <br />chen texanus, (b) Gila elegans, (c) Ptychocheilus Lucius, and (d) <br />Gila cypha. Fish lengths are not to scale, however drawings <br />were made from actual fish specimens courtesy of Jim Beard, <br />U.S. Fish & Wildlife Service. <br />inhabit eddies, deep runs, and slack waters (re- <br />viewed by U.S. Fish & Wildlife Service 1990, 1998, <br />unpublished observations). These are large, but <br />quiet habitats where adaptations to fast flows are <br />not required. Conversely, related native cyprinids <br />(Gila elegans and G. robusta) and catostomids <br />(Catostomus latipinnis and C. discobolus) that <br />prefer fast flowing habitats do not have such large <br />humps. Body shapes of these species suggest low <br />pressure drag, which would be more adaptive to <br />high mobility and fast flows (Webb 1984). Such <br />body shapes tend to have the transitional point <br />between laminar and turbulent flow located far <br />back from the leading edge. In contrast, the tran- <br />sitional point in G. cypha and X. texanus occurs on <br />the anterior leading portion of the body, a mor- <br />phology not conducive to reducing drag (i.e., the