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Hydroclimate Mediates Effects of a Keystone Species in a Coldwater Reservoir Brett M

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Hydroclimate Mediates Effects of a Keystone Species in a Coldwater Reservoir Brett M This article was downloaded by: [Patrick Martinez] On: 27 March 2012, At: 10:25 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK Lake and Reservoir Management Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/ulrm20 Hydroclimate mediates effects of a keystone species in a coldwater reservoir Brett M. Johnson a & Patrick J. Martinez b c a Department of Fish, Wildlife and Conservation Biology, 1474 Campus Delivery, Colorado State University, Fort Collins, CO, 80523 b Colorado Division of Wildlife, 711 Independence Ave., Grand Junction, CO, 81505 c US Fish and Wildlife Service, 764 Horizon Drive, Grand Junction, CO, 81506-8721 Available online: 19 Mar 2012 To cite this article: Brett M. Johnson & Patrick J. Martinez (2012): Hydroclimate mediates effects of a keystone species in a coldwater reservoir, Lake and Reservoir Management, 28:1, 70-83 To link to this article: http://dx.doi.org/10.1080/07438141.2012.658492 PLEASE SCROLL DOWN FOR ARTICLE Full terms and conditions of use: http://www.tandfonline.com/page/terms-and-conditions This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae, and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand, or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Lake and Reservoir Management, 28:70–83, 2012 C Copyright by the North American Lake Management Society 2012 ISSN: 0743-8141 print / 1040-2381 online DOI: 10.1080/07438141.2012.658492 Hydroclimate mediates effects of a keystone species in a coldwater reservoir Brett M. Johnson1,∗ and Patrick J. Martinez2,3 1Department of Fish, Wildlife and Conservation Biology, 1474 Campus Delivery, Colorado State University, Fort Collins, CO 80523 2Colorado Division of Wildlife, 711 Independence Ave., Grand Junction, CO 81505 3Current address: US Fish and Wildlife Service, 764 Horizon Drive, Grand Junction, CO 81506-8721 Abstract Johnson, BM and PJ Martinez 2012. Hydroclimate mediates effects of a keystone species in a coldwater reservoir. Lake Reserv Manage. 28:70–83. Many reservoirs exhibit interannual variations in water content resulting from climatic variability. Hydroclima- tological factors may also affect physicochemical conditions such as thermal stratification and dissolved oxygen concentrations, which in turn have important implications for reservoir biota. We investigated the interplay among water budget, climate, and food web interactions involving a keystone planktivore, Mysis diluviana, occurring at Lake Granby, Colorado, during 1954–2009. A popular kokanee (Oncorhynchus nerka) fishery has been maintained at this 2938 ha montane reservoir, but survival and growth of kokanee has varied widely, with spawning adults ranging in size from 272 to 449 mm total length and population size varying by an order of magnitude (21–269 fish/ha). The kokanee population was strongly affected by Mysis, through competition for food (Daphnia) and because Mysis probably enhanced lake trout recruitment and predation on kokanee. Physicochemical conditions in the reservoir, driven by hydroclimatologic factors, mediated the effects of Mysis on Daphnia, kokanee, and presumably lake trout by controlling habitat suitability, distribution, and survival of Mysis. Effective fishery management in reservoirs requires an improved understanding of the complex interactions among hydroclimate, reservoir conditions, and food web dynamics. Key words: Daphnia, food web, kokanee, Mysis, reservoir management Two intriguing characteristics of reservoirs distinguish them Kokanee (Oncorhynchus nerka) is a popular sport fish in Downloaded by [Patrick Martinez] at 10:25 27 March 2012 from many natural lakes: (1) their inherent variability, and coldwater reservoirs throughout the western United States (2) the degree to which they can respond to anthropogenic and Canada (Rieman and Myers 1992, Martinez et al. 2009). factors (Thornton et al. 1990). Reservoirs in the arid west- These fish are proficient zooplanktivores, efficiently chan- ern United States are exemplary, with annual water level neling a reservoir’s pelagic production into desirable fish fluctuations that can exceed 20 m even in large (≥103 ha) biomass (Koski and Johnson 2002). The introduction of reservoirs (Johnson and Goettl 1999, Baldwin et al. 2003), opossum shrimp (Mysis diluviana,formerlyM. relicta; Rud- and interannual variation in storage is closely tied to climatic stam and Johannsson 2009) into many western reservoirs conditions. Sport fisheries are often subsidized or sustained in the 1970s and 1980s proved detrimental to kokanee, by fish stocking, and fishery quality varies with stocking suc- as Mysis displayed effective predator avoidance behavior cess, which can be linked to reservoir conditions (Bowles et and became a competitor for food rather than a supple- al. 1991, Martinez and Wiltzius 1995, Johnson and Goettl mental food item for kokanee (Nesler and Bergersen 1991). 1999). Effective management of reservoir fisheries requires Depending on thermal conditions, Mysis can contribute to an understanding of the relative roles of anthropogenic and the decline of kokanee populations through “middle out” natural factors in ecosystem dynamics. control (control of food web from an intermediate trophic level; Stein et al. 1995) of food web interactions (Chipps and Bennett 2000). This disruption of a trophic cascade has Mysis outcompeting kokanee for zooplankton prey, but ∗Corresponding author: [email protected] Mysis are undeterred by depletion of their preferred prey 70 Keystone species and hydroclimate because they are omnivorous, subsisting on phytoplank- in 1949 primarily for irrigation; hydrology of the system is ton and detritus when zooplankton are scarce (Northcote complex. Winter snowpack is the primary source of water 1991, Chipps and Bennett 2000). Further, abundant Mysis supply to the reservoir (Matter et al. 2010). Water is also populations seem to foster lake trout (Salvelinus namaycush) pumped into Lake Granby from 2 downstream reservoirs, recruitment by alleviating a trophic bottleneck for juvenile Willow Creek and Windy Gap. Water is released from Lake lake trout (Bowles et al. 1991, Martinez et al. 2009, Ellis et Granby into the Colorado River via a hypolimnetic outlet on al. 2011). Few other sport fish in western lakes and reservoirs the dam but also via a pump that shuttles hypolimnetic water are effective predators of Mysis. Unfortunately for kokanee, upstream into Shadow Mountain Reservoir and eventually as lake trout grow they become piscivorous, with salmonids through the Adams Tunnel across the continental divide to making up the bulk of the diet when available (Johnson and Colorado’s more populous Front Range. Annual drawdown Martinez 2000, Ruyzicki et al. 2003, Martinez et al. 2009). of the reservoir is about 10 m, from an annual low in March Many formerly robust kokanee fisheries across the western and annual high in July. Average content in July is about United States have collapsed, succumbing to the combined 80% of capacity but interannual variation is high, depend- effects of Mysis and lake trout (Martinez et al. 2009). Fishery ing on inflow and user demand. The lowest annual maximum managers striving to restore kokanee in western reservoirs surface elevation was about 19 m below full level (50.2% of require an understanding of how these complex food web in- capacity; Jul 1956), and the lowest annual minimum was 35 teractions involving Mysis (and lake trout) may be mediated m below full level (16.2% of capacity; Apr 1978). In addi- by environmental conditions. tion to Lake Granby discharge, Windy Gap Reservoir also receives water treatment plant effluent. Thus, water from It has been proposed that reduced reservoir content allows Windy Gap Reservoir pumped back into Lake Granby sup- faster spring warming and a warmer epilimnion, which pro- plements Lake Granby’s nutrient supply (Lieberman 2008). vides a thermal refuge for Daphnia and alleviates compe- tition between kokanee and Mysis (Martinez and Wiltzius The food web is typical of many coldwater, montane reser- 1995). This thermal relationship and its mediating effect on voirs of the intermountain west. Water level fluctuations Mysis impacts to zooplankton have been similarly proposed preclude a productive littoral zone. Most consumer biomass for some lakes (Schoen 2007, Koksvik et al. 2009). An ex- derives from pelagic primary production and detritivory in citing implication of the hypothesis for reservoirs suggests the benthos (Johnson et al. 2002). The system is mesotrophic that dam operations offer a means to manipulate entire food (USEPA 1977) with a trophic state index of about 45 webs to channel energy flow away from Mysis and their and mean summer chlorophyll a concentration of about beneficiaries, such as lake trout, and foster increased pro- 4.0 µg/L (Johnson and Martinez
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