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RECOVERY OF THE FOUR BIG-RIVER FISHES OF THE UPPER COLORADO RIVER BASIN
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Recovery of the Four Big-River Fishes of the Upper Colorado River Basin
Christi Meismer
Western Governor 's University
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In the upper Colorado River basin, four species of fish have been listed as federally endangered. In 1988, the Upper Colorado River Endangered Fish Recovery Program
(Program) was created through a distinctive agreement between a …show more content…
suite of entities comprising local, state, and federal agencies, water and power agencies, and environmental and tribal interests (Carlson & Muth, 1993; United States Department of the Interior [USDOI], 1987). The Program was established to recover the endangered fishes while allowing for water development to proceed according to federal and state laws. To achieve the recovery goals established by the U.S. Fish and Wildlife Service, the
Program has embarked on a series of recovery plans focused on three primary activities.
The combination of these plans is considerably different from other recovery efforts that focus on single species or activity to recover a species. Often times, these singular efforts are not successful, but efforts to focus more on benefiting the overall ecosystem are successful. The first action that the Program has undertaken is the manipulation of streamflows from reservoirs to more closely mimic historical conditions to which the endangered fishes were accustomed. Secondly, the Program has worked to reduce the negative impacts of competition and predation that nonnative fish species are having on the native fish. Finally in order to sustain and increase endangered fish populations, hatchery-reared fish are being stocked into the river. Research suggests that without flow
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Colorado River basin will not be de-listed (recovered). In this paper, I will provide an introduction as to why the four big-river fishes of the Colorado River basin are endangered and information on delisting of a species.
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Background/History
The Colorado River is one of the single most important bodies of water in the United
States. Its life-giving water is the only single reason why the western United States has been able to be settled, farmed, and ultimately developed, giving way to the great metropolises such as Phoenix, Los Angeles, Las Vegas and many others. The Colorado
River today supplies more water for consumptive use than any other river in the United
States (Carlson & Muth. 1989). However, the development of this great resource for mankind has put other species on the brink of extinction. In the history of the Colorado
River, there were initially 54 species of fish that were native. Within those 54 species,
83% of those species were endemic (only found in that area) to the Colorado River basin
(Carlson & Muth, 1989). Four of these species, the big-river species, Colorado pikeminnow (Ptychocheilus lucius), razorback sucker (Xyrauchen texanus), humpback chub (Gila cypha), and bonytail (Gila elegans), found in both the Upper and Lower
Colorado River basin are federally listed as endangered. Endangered species are species that are in imminent danger of becoming extinct. With the advent of the Endangered
Species Act of 1973 (ESA) and the establishment of the Upper Colorado River
Endangered Fish Recovery Program, efforts were being made to pull these four species once known as "trash" or "coarse" fish (Quartarone, 1995) from the chasm of extinction.
Recovery of the population of an endangered species is a long process from listing, to implementing recovery actions and finally the "downlisting" or "de-listing" of the species, or its partial recovery or removal from the federal list of endangered species.
The reasons for the decline of the big-river fishes of the upper Colorado River basin are many and explicably linked to the growth and development of the communities of the western United States. The primary issue that led to the decline of these species
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was the regulation of the river through the construction of large mainstem dams
(impoundments). The impoundment of the river into reservoirs drastically altered the biological and hydrologic conditions to which these species had evolved.
The dams created large, deep reservoirs upstream, that released water for hydropower that was consistently cold, moving fish from areas near the dams as well as negatively affecting native fish reproduction by extending hatching periods, reducing growth, and making native fish larvae (young fish) more susceptible to predation by
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The dams and their operations also changed the types of fish that were found in the river, as the conditions (temperatures, sediment loads and habitat) changed significantly enough to allow nonnative species that were either purposely planted or accidentally released to establish and thrive in basin. These species include small minnow species (red shiner, fathead minnow, sand shiner, redside shiner) to large predatory sport fish
(northern pike, channel catfish, smallmouth bass, walleye). From the smallest to the largest nonnative species, they have the ability to prey upon the native fishes. The smaller species prey upon native fish larvae in backwater habitats as well as competing with them for space and food. The larger species prey upon the adult native fishes, taking away the ability of the species to effectively reproduce and recruit.
The four big-river fishes of the Colorado River basin (Colorado pikeminnow, razorback sucker, humpback chub, and bonytail) are jeopardized by large mainstem dams, water diversion and nonnative fish species. The changes in the Colorado River basin have altered the river from a stream that flowed with enough force to carve canyons out of bedrock into a harnessed stream serving the people who live along its length (Ono 1983).
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The four big-river species had adapted over millions of years to the difficult conditions imposed upon them by the Colorado River, extremes of streamflows, temperatures, sediment, and geology. Each species is unique and faces individual challenges in recovery. The Colorado pikeminnow is largest of the four big-river fishes and also the top native predator in the system. Once found throughout all of the Colorado River basin, from Wyoming downstream into Mexico, the pikeminnow migrated over 200 miles to and from spawning areas (United States Fish and Wildlife Service [USFWS], 2002a) .
Pikeminnow were probably the species most affected by the presence of the dams
(Flaming Gorge, Hoover, and Glen Canyon) as their life history is so significantly tied to their spawning migrations. With the dams, not only could they not move to their spawning grounds, but their offspring were washed into sport fish infested reservoirs if the eggs managed to hatch in cold clear waters. Pikeminnow also do not reproduce until they are approximately four years old. Historically, it would not be necessary for pikeminnow to successfully reproduce each year for the species to survive, however, with the changes to the system, now annual reproductive success is critical, and this life history trait is inhibiting that success. Razorback sucker are the most distinct of the four big-river fishes. The razorback gets its name from the distinctive nuchal hump immediately behind its head that serves to help it stay positioned in the flows of the current. Razorback suckers are also one of the two species that have persisted in both the upper and lower basins. Adults grow to a size of three feet (one meter) and have been known to spawn over submerged gravel bars in the river and rocky shoals in lower basin reservoirs (USFWS, 2002b). Razorbacks have been significantly impacted by the
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introduction of nonnative fishes as they specifically prey upon and compete with larval razorback suckers. Another threat to the razorback is the presence of other closely related suckers (both native and nonnative) that provide the possibility for hybridization.
Humpback chub is the other species still found in both the upper and lower basins. As with the razorback sucker, the humpback chub has a prominent nuchal hump, but a hump as opposed to the sharp-edged shape of the "razor"back sucker 's ridge. Only six extant populations exist, all found well within deep, swift canyon reaches of main stem and large tributaries of the Colorado River basin (USFWS, 2002c).
Bonytail, are arguably the most critically endangered, but least understood of the four fish species. Not listed as endangered until 1980, the bonytail is the only one of the four species to be primarily found only in the lower Colorado River basin once recovery was undertaken. Due to the lateness of their discovery, little is known of their preferred habitats outside of historical references (USFWS, 2002d). They were accorded their name due to the pencil-thin caudal peduncle, an adaptation to dealing with the high streamflows of the basin.
The effort to delist the four big-river fish species of the upper Colorado River basin has been led by the Upper Colorado River Endangered Fish Recovery Program. The
Program has relied upon flow manipulation, nonnative fish control and stocking of hatchery reared fish in their efforts to recovery these species. If these species are to be pulled from the brink of extinction, these are the means that must occur.
Flow Manipulation
Studies and adaptive management have indicated that flow manipulation is the most critical of the three main efforts that must occur to recover the endangered fish
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species in the upper Colorado River basin because water drives everything as it relates to fish. While the most important, flow manipulation is also the most difficult due to the number of laws affecting water rights and deliveries. The Colorado River Compact of
1922 is one of the most significant of those, detailing how water is divided between the upper and lower Colorado River basin and between the various states. Water is further divided between agriculture, municipal, and energy interests. The needs of the native fishes ultimately were an afterthought to the needs of the people. When it comes to the necessary manipulation of flows, this can create significant issues (water needs, flood control, etc). Thirteen years after the establishment of the Program, Muth et al. (2000) identified a series of recommendations aimed at altering the flows out of Flaming Gorge
Reservoir, Green River, Wyoming for the benefit of the native fish, while meeting the needs of the communities. The recommendations are intended to create variation both within years as well as between years to create physiographic and geologic changes in the
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In addition to the flow recommendations on the Green River provided by Muth et al.
(2000), McAda (2003) provided flow recommendations on the Colorado and Gunnison rivers. Similar to those on the Green River, the recommendations were based upon water year. As with Muth, McAda (2003) identified the effects that these flow recommendations would have on the hydrology of the river and ultimately the habitat required by the endangered fishes at different life stages.
Modde and Keleher (2003) proposed similar recommendations for the Duchesne
River, a tributary to the Green River in Utah, although the Duchesne River flows are intended only to provide benefit for adult Colorado pikeminnow. Sampling has shown
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that the Duchesne River is primarily used by in the spring and summer months, particularly by Colorado pikeminnow for temperature preference and to escape the high flows of the Green River while foraging. The move by the Program to create and implement the series of flow recommendations on every major river within the basin illustrates the importance of the effort to the endangered fishes as well as the willingness of the participants within the Program to work towards the recovery of these fish species. Flow manipulation will also have effects on the activities to reduce negative interactions between the endangered fish and nonnative fish species.
Nonnative Removal
The second major endeavor of that must happen for the endangered fishes of the
Colorado River basin to be delisted is to reduce if not eliminate negative interactions between the native fish and nonnative species that have been introduced into the basin
(USFWS 2002a, 2002b, 2002c, 2002d; Lentsch, Muth, Thompson, Hoskins, &
Crowl,1996; Tyus & Stauffer, 1996). Research suggests that native fish recovery in the
Colorado River basin can be achieved by managing the interactions with nonnative fish.
Currently nonnative species comprise the majority of the fish biomass in the basin. The impacts that these nonnative species have can be observed as competition for habitat and food as well as predation on all life stages of the endangered fishes (Lentsch et al., 1996).
Efforts to reduce the impacts of these species included restricting stocking of nonnative fish, screening outlets from recreational reservoirs, and attempts to physically remove the nonnative fish from the rivers (Nesler, 1996; Martinez, 2004; Fuller, 2009; Bestgen et al.,
2009; Trammell et al., 2005). The nonnative fishes that have become established within the basin can typically be separated into two different groups, recreational sport fish and
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non-sport fish introductions. The sport fish introductions have historically and typically been made by official agencies responsible for creating recreational fishing opportunities.
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Following the closure of Flaming Gorge Dam in 1969, these agencies poisoned the Green
River below the dam to remove the native fishes to allow for the establishment of a tailwater trout fishery. Not only did this activity directly affect the populations of native species, but it also affected populations long term through predation and competition with the nonnative species introduced post-poisoning. Additional nonnative sport fish such as largemouth bass and green sunfish have been introduced to the many reservoirs and offstream ponds within the Colorado River basin. Many of these facilities, when full, will release water and nonnative species into the rivers where they interact with native fish.
Controlling these introductions has become more of a focus of the Program over the years. In 1996 three state wildlife agencies (Colorado, Utah, and Wyoming) and the U.S.
Fish and Wildlife Service completed a report (USFWS 1996) that outlined procedures for stocking nonnative fish species in the upper Colorado River basin. While these procedures did help to address some of the issues associated with nonnative fishes in the basin by restricting new species beyond what was already present, they did not address the escapement from reservoirs and ultimately establishment in the river. Martinez and
Nibbelink (2004) were the first to evaluate the effects of the stocking plans, with specific emphasis within the State of Colorado. They determined that, unfortunately, the ability to track stocking events made it nearly impossible to determine exactly what effect the stocking regulations had on endangered fish recovery. Efforts in the interim have attempted to place nets or other screening mechanisms at outlets from reservoirs with varying results along the Colorado and Duchesne rivers (Martinez, 2004;
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USFWS, 2011). Martinez (2004) showed that in most cases (2 of 54 ponds) where largemouth bass were eliminated from a pond, they did not re-establish during the course of the study. However, they further determined that no evidence of their removal efforts could be observed in the river due to existing fish as well as the inability to effect removal on all ponds in the area. In the Duchesne River, a tributary to the Green River, a screen on Crystal Reservoir has prevented escapement of nonnative fishes (USFWS, 2011). In addition to managing the stocking and escapement of fish from reservoirs, the Program has placed significant time, effort and money into physically removing nonnative fishes from the river.
Efforts to remove nonnative predatory fishes from the upper basin rivers have been varied from angling (Fuller, 2009) to electrofishing efforts (Hawkins, Walford, & Hill,
2009). The efforts by Hawkins et al. (2009) to remove smallmouth bass from the Yampa
River using electrofishing illustrated the benefit of removal activities. Hawkins et al. saw a decrease in smallmouth bass numbers that they attributed to their efforts, but in turn determined that immigration of fish into the area and reproduction likely reduced the overall effectiveness of the removal efforts.
Fuller (2009) worked to remove nonnative channel catfish and smallmouth bass from the Yampa River through angling and electrofishing from 2001-2006. Angling efforts for channel catfish were discontinued after 2003 because smallmouth bass populations were increasing and electrofishing is much more effective in their capture and removal, a good
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populations to a point that they would not impact native fish recovery. However, while not reaching the goal, the efforts were trending in the proper direction, but would require more effort to achieve exploitation of the populations. Bestgen et al. (2009) provided an accompanying analysis of Fuller evaluating the native fish response to nonnative fish removal. These researchers determined that native fish populations did not respond to the removal of nonnative fish for a few different reasons. Bestgen et al. (2009) hypothesized that native fish numbers did not increase due to an insufficient number of nonnative fish being removed, changes in streamflows and water temperatures changed as a result of drought to conditions favoring the predatory fishes, and there may have been a lack of reproductively viable native fishes to respond to the decreased nonnative populations.
Ultimately they determined that additional efforts in both fish removal and monitoring of native fish response will be necessary.
While Fuller (2009) and Martinez (2004) were removing sport fish, Trammell et al.
(2005) detailed the efforts over three years to remove the small-bodied nonnative minnows (red shiner, fathead minnow, and sand shiner) from the Green and Colorado rivers as part of one of these studies. While these species are ultimately much smaller than the natives, adults are present in nursery habitats and prey upon native fish larvae
(young fish) as they drift into the nursery habitats. Nonnative fish in these areas outnumber natives sometimes thousands to one. Removal efforts were conducted through seining (pulling a net through the water) in backwater habitats where these species were
predominant.
The researchers found that while hundreds of thousands of these fish were removed, no reach scale benefits were found for native fish (Trammell et al. 2005).
However, one of the beneficial items from this research is that it could be
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possible to remove nonnatives just prior to the arrival of drifting native larvae. This information coupled with the effects of temperatures on growth identified by Bestgen
(2008), could allow native larvae to reach sizes where they will be safe from predation and thus survive at higher rates. In other areas of the basin, Utah Division of Wildlife
Resources (2010) biologists, found the highest numbers of young Colorado pikeminnow since 1991, likely a result of nonnative removal efforts. The ability of all life stages of native fishes to survive encounters with nonnative species will be important to overall survival, but also important to how stocking of hatchery-reared fish is managed.
Stocking of Hatchery Reared Fish
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Research suggests that unless hatchery-reared fish are stocked into the river the endangered fishes of the upper Colorado River basin will not be recovered. Stocking of fish to enhance low population numbers has been occurring for endangered species recovery efforts for many years. Due to the long life spans of the endangered Colorado
River fishes along with the fact that they do not reach reproductive maturity for a number of years, stocking has been important to initially stabilize populations and then to augment the wild populations. Currently the Program has propagation capabilities at four different facilities (Dexter National Fish Hatchery and Technology Center, 24 Road Hatchery,
Wahweap State Fish Hatchery, and the John W. Mumma Fish Hatchery). These facilities provide thousands of individual fish to Program managers to enhance existing populations. In 2003, Nesler et al. compared the Colorado, Utah, and Wyoming state stocking plans for the four big river fishes of the Colorado River basin. During that evaluation, they determined that the state plans were inconsistent with regards to numbers to stock, age of adult fish, numbers of adult age classes, and number of years to stock.
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To solve these issues, the Program decided to integrate all of the stocking plans into one single plan. The final numbers identified in the integrated plan were designed to meet the goals of the individual species recovery plans (USFWS 2002a, 2002b, 2002c, 2002d).
While hatcheries can consistently provide fish for stocking into the rivers, stocking will be most effective with continued evaluation of successes. As biologists monitor the status of populations, they will in turn discuss with hatcheries and others to determine the best uses of each year 's output of fish. Hatcheries each year attempt to produce enough fish to meet the stocking goals.
Often, the monitoring efforts must not only look at simple hatchery production, but how that production is surviving in the river. The recovery goals for razorback sucker require that two separate populations represented by 5,800 individuals each will be necessary for downlisting or delisting to occur. Zelasko, Bestgen, & White, (2010) evaluated the effectiveness of the stocking plan for razorback sucker. The authors determined that stocking success was increased by not stocking razorback sucker during the summer. They also determined that stocking larger fish into the river enhanced survival but recommended that a cost benefit analysis be conducted to determine if producing fewer larger fish is more expensive than producing more, smaller fish for introduction. A few excellent examples of stocking success have been found in the White
River (Loomis, 2011) and also in the San Juan River. In the White River, razorback sucker larvae from previously stocked fish have been found in the river for the first time.
In the San Juan River, stocking of both Colorado pikeminnow and razorback sucker have been successful enough that both species have reproduced and now there are
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growing populations. Ultimately, continually refining the production from hatcheries has the best potential to significantly help in the recovery process.
Conclusion
In conclusion, since the western United States was settled, mankind 's dependence on the Colorado River increased steadily. Water for agriculture, culinary, and municipal uses were removed from the river. In order for these needs to be fulfilled, large dams were constructed on the river, thereby altering the hydrologic characteristics of the river basin and impacting the native fishes that adapted to the extreme conditions of the basin. The regulation of the river created changes that reduced the high spring flows, lowered water temperatures, and created barriers to spawning migrations of the endangered fishes. The effects of these changes on the native fish were staggering. Nonnative fish were stocked into newly created reservoirs, where they escaped and were able to establish and thrive in the newly regulated river system. The flows released from reservoirs, cold and clear result in lower growth rates for native fish that make them more susceptible to predation by the nonnative fish, ultimately placing them in danger of extinction. These issues have pushed the four big river fishes of the Colorado River to the brink of extinction, requiring human intervention to insure that that extinction does not occur.
The effort to recover the endangered fishes of the upper Colorado River basin has been led by the Upper Colorado River Endangered Fish Recovery Program (Program).
Research conducted as part of the Program has shown that by focusing on the manipulation of flows from reservoirs, controlling nonnative fishes, and stocking of hatchery-reared native fishes will allow the fishes of the Colorado River to be de-listed
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(recovered). The manipulation of flows from dams will reverse some of the hydrologic impacts, re-creating portions the historic extremes of flow found in the basin, and enhance conditions that will likely favor the native fish more than the altered conditions.
The establishment of nonnative fish throughout the Colorado River basin likely represents the largest threat to the endangered fishes. A diverse suite of nonnative fish, compete with and prey upon the native fish, reducing their ability to thrive. Efforts to control these fish, include management of future introductions, attempts to restrict escapement from reservoirs, and physical removal of these species. These efforts are yielding positive results, but gains are measured in small steps and require constant adaptive management.
The use of hatcheries to provide large numbers of endangered fish is a time-tested means of stabilizing and increasing populations of fish. Stocking of the endangered fishes into the Colorado River basin has been ongoing for a number of years and fish stocked are now beginning to return and spawn, creating wild fish that will continue to enhance population numbers. All of these efforts must occur for these fishes to be de-listed.
Currently, gains in the native populations are moving slowly towards downlisting and delisting as a result of the focused efforts to address flow manipulation, nonnative species control and stocking of hatchery-reared fishes.
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Region (6), Denver, Colorado.
---------. (2011). Nonnative Fish Management Questions and Answers – 2011 (Utah). http://www.coloradoriverrecovery.org/events-news/news/NNF-Q&A-UT-2011.pdf. Zelasko, K., Bestgen, K., & White, G. (2010). Survival rates and movement of hatcheryreared razorback suckers in the Upper Colorado River basin, Utah and Colorado.
Transactions of the American Fisheries Society 139, 1478–1499.
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RECOVERY OF THE FOUR BIG-RIVER FISHES OF THE UPPER COLORADO RIVER BASIN
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Recovery of the Four Big-River Fishes of the Upper Colorado River Basin
Christi Meismer
Western Governor 's University
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In the upper Colorado River basin, four species of fish have been listed as federally endangered. In 1988, the Upper Colorado River Endangered Fish Recovery Program
(Program) was created through a distinctive agreement between a …show more content…
suite of entities comprising local, state, and federal agencies, water and power agencies, and environmental and tribal interests (Carlson & Muth, 1993; United States Department of the Interior [USDOI], 1987). The Program was established to recover the endangered fishes while allowing for water development to proceed according to federal and state laws. To achieve the recovery goals established by the U.S. Fish and Wildlife Service, the
Program has embarked on a series of recovery plans focused on three primary activities.
The combination of these plans is considerably different from other recovery efforts that focus on single species or activity to recover a species. Often times, these singular efforts are not successful, but efforts to focus more on benefiting the overall ecosystem are successful. The first action that the Program has undertaken is the manipulation of streamflows from reservoirs to more closely mimic historical conditions to which the endangered fishes were accustomed. Secondly, the Program has worked to reduce the negative impacts of competition and predation that nonnative fish species are having on the native fish. Finally in order to sustain and increase endangered fish populations, hatchery-reared fish are being stocked into the river. Research suggests that without flow
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Colorado River basin will not be de-listed (recovered). In this paper, I will provide an introduction as to why the four big-river fishes of the Colorado River basin are endangered and information on delisting of a species.
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Background/History
The Colorado River is one of the single most important bodies of water in the United
States. Its life-giving water is the only single reason why the western United States has been able to be settled, farmed, and ultimately developed, giving way to the great metropolises such as Phoenix, Los Angeles, Las Vegas and many others. The Colorado
River today supplies more water for consumptive use than any other river in the United
States (Carlson & Muth. 1989). However, the development of this great resource for mankind has put other species on the brink of extinction. In the history of the Colorado
River, there were initially 54 species of fish that were native. Within those 54 species,
83% of those species were endemic (only found in that area) to the Colorado River basin
(Carlson & Muth, 1989). Four of these species, the big-river species, Colorado pikeminnow (Ptychocheilus lucius), razorback sucker (Xyrauchen texanus), humpback chub (Gila cypha), and bonytail (Gila elegans), found in both the Upper and Lower
Colorado River basin are federally listed as endangered. Endangered species are species that are in imminent danger of becoming extinct. With the advent of the Endangered
Species Act of 1973 (ESA) and the establishment of the Upper Colorado River
Endangered Fish Recovery Program, efforts were being made to pull these four species once known as "trash" or "coarse" fish (Quartarone, 1995) from the chasm of extinction.
Recovery of the population of an endangered species is a long process from listing, to implementing recovery actions and finally the "downlisting" or "de-listing" of the species, or its partial recovery or removal from the federal list of endangered species.
The reasons for the decline of the big-river fishes of the upper Colorado River basin are many and explicably linked to the growth and development of the communities of the western United States. The primary issue that led to the decline of these species
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was the regulation of the river through the construction of large mainstem dams
(impoundments). The impoundment of the river into reservoirs drastically altered the biological and hydrologic conditions to which these species had evolved.
The dams created large, deep reservoirs upstream, that released water for hydropower that was consistently cold, moving fish from areas near the dams as well as negatively affecting native fish reproduction by extending hatching periods, reducing growth, and making native fish larvae (young fish) more susceptible to predation by
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The dams and their operations also changed the types of fish that were found in the river, as the conditions (temperatures, sediment loads and habitat) changed significantly enough to allow nonnative species that were either purposely planted or accidentally released to establish and thrive in basin. These species include small minnow species (red shiner, fathead minnow, sand shiner, redside shiner) to large predatory sport fish
(northern pike, channel catfish, smallmouth bass, walleye). From the smallest to the largest nonnative species, they have the ability to prey upon the native fishes. The smaller species prey upon native fish larvae in backwater habitats as well as competing with them for space and food. The larger species prey upon the adult native fishes, taking away the ability of the species to effectively reproduce and recruit.
The four big-river fishes of the Colorado River basin (Colorado pikeminnow, razorback sucker, humpback chub, and bonytail) are jeopardized by large mainstem dams, water diversion and nonnative fish species. The changes in the Colorado River basin have altered the river from a stream that flowed with enough force to carve canyons out of bedrock into a harnessed stream serving the people who live along its length (Ono 1983).
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The four big-river species had adapted over millions of years to the difficult conditions imposed upon them by the Colorado River, extremes of streamflows, temperatures, sediment, and geology. Each species is unique and faces individual challenges in recovery. The Colorado pikeminnow is largest of the four big-river fishes and also the top native predator in the system. Once found throughout all of the Colorado River basin, from Wyoming downstream into Mexico, the pikeminnow migrated over 200 miles to and from spawning areas (United States Fish and Wildlife Service [USFWS], 2002a) .
Pikeminnow were probably the species most affected by the presence of the dams
(Flaming Gorge, Hoover, and Glen Canyon) as their life history is so significantly tied to their spawning migrations. With the dams, not only could they not move to their spawning grounds, but their offspring were washed into sport fish infested reservoirs if the eggs managed to hatch in cold clear waters. Pikeminnow also do not reproduce until they are approximately four years old. Historically, it would not be necessary for pikeminnow to successfully reproduce each year for the species to survive, however, with the changes to the system, now annual reproductive success is critical, and this life history trait is inhibiting that success. Razorback sucker are the most distinct of the four big-river fishes. The razorback gets its name from the distinctive nuchal hump immediately behind its head that serves to help it stay positioned in the flows of the current. Razorback suckers are also one of the two species that have persisted in both the upper and lower basins. Adults grow to a size of three feet (one meter) and have been known to spawn over submerged gravel bars in the river and rocky shoals in lower basin reservoirs (USFWS, 2002b). Razorbacks have been significantly impacted by the
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introduction of nonnative fishes as they specifically prey upon and compete with larval razorback suckers. Another threat to the razorback is the presence of other closely related suckers (both native and nonnative) that provide the possibility for hybridization.
Humpback chub is the other species still found in both the upper and lower basins. As with the razorback sucker, the humpback chub has a prominent nuchal hump, but a hump as opposed to the sharp-edged shape of the "razor"back sucker 's ridge. Only six extant populations exist, all found well within deep, swift canyon reaches of main stem and large tributaries of the Colorado River basin (USFWS, 2002c).
Bonytail, are arguably the most critically endangered, but least understood of the four fish species. Not listed as endangered until 1980, the bonytail is the only one of the four species to be primarily found only in the lower Colorado River basin once recovery was undertaken. Due to the lateness of their discovery, little is known of their preferred habitats outside of historical references (USFWS, 2002d). They were accorded their name due to the pencil-thin caudal peduncle, an adaptation to dealing with the high streamflows of the basin.
The effort to delist the four big-river fish species of the upper Colorado River basin has been led by the Upper Colorado River Endangered Fish Recovery Program. The
Program has relied upon flow manipulation, nonnative fish control and stocking of hatchery reared fish in their efforts to recovery these species. If these species are to be pulled from the brink of extinction, these are the means that must occur.
Flow Manipulation
Studies and adaptive management have indicated that flow manipulation is the most critical of the three main efforts that must occur to recover the endangered fish
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species in the upper Colorado River basin because water drives everything as it relates to fish. While the most important, flow manipulation is also the most difficult due to the number of laws affecting water rights and deliveries. The Colorado River Compact of
1922 is one of the most significant of those, detailing how water is divided between the upper and lower Colorado River basin and between the various states. Water is further divided between agriculture, municipal, and energy interests. The needs of the native fishes ultimately were an afterthought to the needs of the people. When it comes to the necessary manipulation of flows, this can create significant issues (water needs, flood control, etc). Thirteen years after the establishment of the Program, Muth et al. (2000) identified a series of recommendations aimed at altering the flows out of Flaming Gorge
Reservoir, Green River, Wyoming for the benefit of the native fish, while meeting the needs of the communities. The recommendations are intended to create variation both within years as well as between years to create physiographic and geologic changes in the
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In addition to the flow recommendations on the Green River provided by Muth et al.
(2000), McAda (2003) provided flow recommendations on the Colorado and Gunnison rivers. Similar to those on the Green River, the recommendations were based upon water year. As with Muth, McAda (2003) identified the effects that these flow recommendations would have on the hydrology of the river and ultimately the habitat required by the endangered fishes at different life stages.
Modde and Keleher (2003) proposed similar recommendations for the Duchesne
River, a tributary to the Green River in Utah, although the Duchesne River flows are intended only to provide benefit for adult Colorado pikeminnow. Sampling has shown
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that the Duchesne River is primarily used by in the spring and summer months, particularly by Colorado pikeminnow for temperature preference and to escape the high flows of the Green River while foraging. The move by the Program to create and implement the series of flow recommendations on every major river within the basin illustrates the importance of the effort to the endangered fishes as well as the willingness of the participants within the Program to work towards the recovery of these fish species. Flow manipulation will also have effects on the activities to reduce negative interactions between the endangered fish and nonnative fish species.
Nonnative Removal
The second major endeavor of that must happen for the endangered fishes of the
Colorado River basin to be delisted is to reduce if not eliminate negative interactions between the native fish and nonnative species that have been introduced into the basin
(USFWS 2002a, 2002b, 2002c, 2002d; Lentsch, Muth, Thompson, Hoskins, &
Crowl,1996; Tyus & Stauffer, 1996). Research suggests that native fish recovery in the
Colorado River basin can be achieved by managing the interactions with nonnative fish.
Currently nonnative species comprise the majority of the fish biomass in the basin. The impacts that these nonnative species have can be observed as competition for habitat and food as well as predation on all life stages of the endangered fishes (Lentsch et al., 1996).
Efforts to reduce the impacts of these species included restricting stocking of nonnative fish, screening outlets from recreational reservoirs, and attempts to physically remove the nonnative fish from the rivers (Nesler, 1996; Martinez, 2004; Fuller, 2009; Bestgen et al.,
2009; Trammell et al., 2005). The nonnative fishes that have become established within the basin can typically be separated into two different groups, recreational sport fish and
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non-sport fish introductions. The sport fish introductions have historically and typically been made by official agencies responsible for creating recreational fishing opportunities.
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Following the closure of Flaming Gorge Dam in 1969, these agencies poisoned the Green
River below the dam to remove the native fishes to allow for the establishment of a tailwater trout fishery. Not only did this activity directly affect the populations of native species, but it also affected populations long term through predation and competition with the nonnative species introduced post-poisoning. Additional nonnative sport fish such as largemouth bass and green sunfish have been introduced to the many reservoirs and offstream ponds within the Colorado River basin. Many of these facilities, when full, will release water and nonnative species into the rivers where they interact with native fish.
Controlling these introductions has become more of a focus of the Program over the years. In 1996 three state wildlife agencies (Colorado, Utah, and Wyoming) and the U.S.
Fish and Wildlife Service completed a report (USFWS 1996) that outlined procedures for stocking nonnative fish species in the upper Colorado River basin. While these procedures did help to address some of the issues associated with nonnative fishes in the basin by restricting new species beyond what was already present, they did not address the escapement from reservoirs and ultimately establishment in the river. Martinez and
Nibbelink (2004) were the first to evaluate the effects of the stocking plans, with specific emphasis within the State of Colorado. They determined that, unfortunately, the ability to track stocking events made it nearly impossible to determine exactly what effect the stocking regulations had on endangered fish recovery. Efforts in the interim have attempted to place nets or other screening mechanisms at outlets from reservoirs with varying results along the Colorado and Duchesne rivers (Martinez, 2004;
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USFWS, 2011). Martinez (2004) showed that in most cases (2 of 54 ponds) where largemouth bass were eliminated from a pond, they did not re-establish during the course of the study. However, they further determined that no evidence of their removal efforts could be observed in the river due to existing fish as well as the inability to effect removal on all ponds in the area. In the Duchesne River, a tributary to the Green River, a screen on Crystal Reservoir has prevented escapement of nonnative fishes (USFWS, 2011). In addition to managing the stocking and escapement of fish from reservoirs, the Program has placed significant time, effort and money into physically removing nonnative fishes from the river.
Efforts to remove nonnative predatory fishes from the upper basin rivers have been varied from angling (Fuller, 2009) to electrofishing efforts (Hawkins, Walford, & Hill,
2009). The efforts by Hawkins et al. (2009) to remove smallmouth bass from the Yampa
River using electrofishing illustrated the benefit of removal activities. Hawkins et al. saw a decrease in smallmouth bass numbers that they attributed to their efforts, but in turn determined that immigration of fish into the area and reproduction likely reduced the overall effectiveness of the removal efforts.
Fuller (2009) worked to remove nonnative channel catfish and smallmouth bass from the Yampa River through angling and electrofishing from 2001-2006. Angling efforts for channel catfish were discontinued after 2003 because smallmouth bass populations were increasing and electrofishing is much more effective in their capture and removal, a good
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populations to a point that they would not impact native fish recovery. However, while not reaching the goal, the efforts were trending in the proper direction, but would require more effort to achieve exploitation of the populations. Bestgen et al. (2009) provided an accompanying analysis of Fuller evaluating the native fish response to nonnative fish removal. These researchers determined that native fish populations did not respond to the removal of nonnative fish for a few different reasons. Bestgen et al. (2009) hypothesized that native fish numbers did not increase due to an insufficient number of nonnative fish being removed, changes in streamflows and water temperatures changed as a result of drought to conditions favoring the predatory fishes, and there may have been a lack of reproductively viable native fishes to respond to the decreased nonnative populations.
Ultimately they determined that additional efforts in both fish removal and monitoring of native fish response will be necessary.
While Fuller (2009) and Martinez (2004) were removing sport fish, Trammell et al.
(2005) detailed the efforts over three years to remove the small-bodied nonnative minnows (red shiner, fathead minnow, and sand shiner) from the Green and Colorado rivers as part of one of these studies. While these species are ultimately much smaller than the natives, adults are present in nursery habitats and prey upon native fish larvae
(young fish) as they drift into the nursery habitats. Nonnative fish in these areas outnumber natives sometimes thousands to one. Removal efforts were conducted through seining (pulling a net through the water) in backwater habitats where these species were
predominant.
The researchers found that while hundreds of thousands of these fish were removed, no reach scale benefits were found for native fish (Trammell et al. 2005).
However, one of the beneficial items from this research is that it could be
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possible to remove nonnatives just prior to the arrival of drifting native larvae. This information coupled with the effects of temperatures on growth identified by Bestgen
(2008), could allow native larvae to reach sizes where they will be safe from predation and thus survive at higher rates. In other areas of the basin, Utah Division of Wildlife
Resources (2010) biologists, found the highest numbers of young Colorado pikeminnow since 1991, likely a result of nonnative removal efforts. The ability of all life stages of native fishes to survive encounters with nonnative species will be important to overall survival, but also important to how stocking of hatchery-reared fish is managed.
Stocking of Hatchery Reared Fish
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Research suggests that unless hatchery-reared fish are stocked into the river the endangered fishes of the upper Colorado River basin will not be recovered. Stocking of fish to enhance low population numbers has been occurring for endangered species recovery efforts for many years. Due to the long life spans of the endangered Colorado
River fishes along with the fact that they do not reach reproductive maturity for a number of years, stocking has been important to initially stabilize populations and then to augment the wild populations. Currently the Program has propagation capabilities at four different facilities (Dexter National Fish Hatchery and Technology Center, 24 Road Hatchery,
Wahweap State Fish Hatchery, and the John W. Mumma Fish Hatchery). These facilities provide thousands of individual fish to Program managers to enhance existing populations. In 2003, Nesler et al. compared the Colorado, Utah, and Wyoming state stocking plans for the four big river fishes of the Colorado River basin. During that evaluation, they determined that the state plans were inconsistent with regards to numbers to stock, age of adult fish, numbers of adult age classes, and number of years to stock.
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To solve these issues, the Program decided to integrate all of the stocking plans into one single plan. The final numbers identified in the integrated plan were designed to meet the goals of the individual species recovery plans (USFWS 2002a, 2002b, 2002c, 2002d).
While hatcheries can consistently provide fish for stocking into the rivers, stocking will be most effective with continued evaluation of successes. As biologists monitor the status of populations, they will in turn discuss with hatcheries and others to determine the best uses of each year 's output of fish. Hatcheries each year attempt to produce enough fish to meet the stocking goals.
Often, the monitoring efforts must not only look at simple hatchery production, but how that production is surviving in the river. The recovery goals for razorback sucker require that two separate populations represented by 5,800 individuals each will be necessary for downlisting or delisting to occur. Zelasko, Bestgen, & White, (2010) evaluated the effectiveness of the stocking plan for razorback sucker. The authors determined that stocking success was increased by not stocking razorback sucker during the summer. They also determined that stocking larger fish into the river enhanced survival but recommended that a cost benefit analysis be conducted to determine if producing fewer larger fish is more expensive than producing more, smaller fish for introduction. A few excellent examples of stocking success have been found in the White
River (Loomis, 2011) and also in the San Juan River. In the White River, razorback sucker larvae from previously stocked fish have been found in the river for the first time.
In the San Juan River, stocking of both Colorado pikeminnow and razorback sucker have been successful enough that both species have reproduced and now there are
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growing populations. Ultimately, continually refining the production from hatcheries has the best potential to significantly help in the recovery process.
Conclusion
In conclusion, since the western United States was settled, mankind 's dependence on the Colorado River increased steadily. Water for agriculture, culinary, and municipal uses were removed from the river. In order for these needs to be fulfilled, large dams were constructed on the river, thereby altering the hydrologic characteristics of the river basin and impacting the native fishes that adapted to the extreme conditions of the basin. The regulation of the river created changes that reduced the high spring flows, lowered water temperatures, and created barriers to spawning migrations of the endangered fishes. The effects of these changes on the native fish were staggering. Nonnative fish were stocked into newly created reservoirs, where they escaped and were able to establish and thrive in the newly regulated river system. The flows released from reservoirs, cold and clear result in lower growth rates for native fish that make them more susceptible to predation by the nonnative fish, ultimately placing them in danger of extinction. These issues have pushed the four big river fishes of the Colorado River to the brink of extinction, requiring human intervention to insure that that extinction does not occur.
The effort to recover the endangered fishes of the upper Colorado River basin has been led by the Upper Colorado River Endangered Fish Recovery Program (Program).
Research conducted as part of the Program has shown that by focusing on the manipulation of flows from reservoirs, controlling nonnative fishes, and stocking of hatchery-reared native fishes will allow the fishes of the Colorado River to be de-listed
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(recovered). The manipulation of flows from dams will reverse some of the hydrologic impacts, re-creating portions the historic extremes of flow found in the basin, and enhance conditions that will likely favor the native fish more than the altered conditions.
The establishment of nonnative fish throughout the Colorado River basin likely represents the largest threat to the endangered fishes. A diverse suite of nonnative fish, compete with and prey upon the native fish, reducing their ability to thrive. Efforts to control these fish, include management of future introductions, attempts to restrict escapement from reservoirs, and physical removal of these species. These efforts are yielding positive results, but gains are measured in small steps and require constant adaptive management.
The use of hatcheries to provide large numbers of endangered fish is a time-tested means of stabilizing and increasing populations of fish. Stocking of the endangered fishes into the Colorado River basin has been ongoing for a number of years and fish stocked are now beginning to return and spawn, creating wild fish that will continue to enhance population numbers. All of these efforts must occur for these fishes to be de-listed.
Currently, gains in the native populations are moving slowly towards downlisting and delisting as a result of the focused efforts to address flow manipulation, nonnative species control and stocking of hatchery-reared fishes.
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