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Shifts in the Physical and Biological Characteristics of Alpine Lakes Along an Elevation Gradient in the Rocky Mountains, USA

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Shifts in the Physical and Biological Characteristics of Alpine Lakes Along an Elevation Gradient in the Rocky Mountains, USA Aquatic Sciences (2020) 82:11 https://doi.org/10.1007/s00027-019-0684-6 Aquatic Sciences RESEARCH ARTICLE The life aquatic in high relief: shifts in the physical and biological characteristics of alpine lakes along an elevation gradient in the Rocky Mountains, USA Kelly A. Loria1 · Diane McKnight2 · Dillon M. Ragar3 · Pieter T. J. Johnson4 Received: 26 March 2019 / Accepted: 28 November 2019 / Published online: 4 December 2019 © Springer Nature Switzerland AG 2019 Abstract Rapidly occurring environmental changes in alpine lakes highlight the importance of better understanding the ecological structure and function associated with these systems. Previous research has identifed how the physical characteristics of lakes change as a function of landscape position, but comparatively little is known about shifts in the biotic community across mountain regions. In 2016, we sampled 19 lakes across an elevation gradient (2480–3550 m a.s.l.) within the Rocky Moun- tains, USA, to evaluate how both the abiotic characteristics of lakes and their planktonic biological communities covaried with elevation. Based on generalized linear mixed models (GLMMs), increases in elevation were associated with decreases in most nutrient concentrations (with the exception of nitrate), dissolved organic carbon, water temperature and lake stratifcation. Conversely, elevation increases were positively related to nitrate concentrations and water clarity. Extending this analysis to the biological community, we found that higher-elevation lakes exhibited lower phytoplankton and zooplankton densities, whereas elevation associated positively with average zooplankton size. Our data are consistent with the hypothesis that the alpine environment acts as a strong niche flter, limiting the quantity and diversity of taxa to groups capable of tolerating the short growing season, high fushing rate, strong variation in interannual precipitation, intense ultraviolet radiation exposure, and lower resource availability associated with such habitats. Keywords Landscape limnology · Mountain lake · Elevation gradient · Freshwater ecology · Zooplankton · Phytoplankton Introduction Electronic supplementary material The online version of this article (https ://doi.org/10.1007/s0002 7-019-0684-6) contains Worldwide, an estimated 20% of lakes are located higher supplementary material, which is available to authorized users. than 1000 m a.s.l. (Verpoorter et al. 2014). Growing evi- * Kelly A. Loria dence suggests that these mountainous, often remote aquatic [email protected] systems are critical to downstream ecosystems and may 1 difer considerably from low-elevation lakes in both their Niwot Ridge Long Term Ecological Research Program, physical and biological properties (Moser et al. 2019). For Institute of Arctic and Alpine Research, University of Colorado, Campus Box 450, Boulder, CO 80309-0450, instance, mountain lakes often function as a hot spot for USA nutrient cycling (Alexander et al. 2007; Brown et al. 2008) 2 Department of Civil, Environmental and Architectural and are a key source of freshwater for humans (Beniston Engineering, Environmental Studies, Hydrological Sciences, 2003; Viviroli et al. 2007). Relative to lakes at lower eleva- Institute of Arctic and Alpine Research, University tions, which have a much longer history of study (Jacobsen of Colorado, Campus Box 450, Boulder, CO 80309-0450, and Dangles 2017), alpine lakes (those above treeline) are USA 3 characterized by sparse surrounding vegetation, extended Boulder Creek Critical Zone Observatory, Institute of Arctic ice-cover (Caine 2002; Hampton et al. 2017), intense and Alpine Research, University of Colorado, Campus Box 450, Boulder, CO 80309-0450, USA spring fushing rates (McGuire et al. 2005; Clow 2010), and high exposure to solar radiation (Blumthaler et al. 1997). 4 Department of Ecology and Evolutionary Biology, University of Colorado, Ramaley N122, CB334, Boulder, CO 80309, The steep topography, abrupt transition in vegetation pat- USA terns, unique basin shapes, dynamic weather patterns, and Vol.:(0123456789)1 3 11 Page 2 of 16 K. A. Loria et al. interannual variation in winter snowpack of alpine environ- and potentially limiting nutrient concentrations (Hansson ments make it probable that lakes will also vary broadly in et al. 2007; Miller and McKnight 2015). For instance, responses to forecasted environmental changes, underscor- in a survey of lakes in the Pyrenees, macroinvertebrate ing the importance of research on the ecology of mountain richness decreased by 57% for lakes above 2800 m a.s.l. lake ecosystems (Kamenik et al. 2001; Kraemer et al. 2015). (De Mendoza and Catalan 2010). Similarly, Lyons and Previous limnological research has emphasized the Vinebrooke (2016) found that zooplankton species rich- importance of landscape position, or the concept of a lake’s ness in Canadian Rockies decreased from an average of hydrologic position within the local to regional fow sys- 10.7 species per lake for lakes at or below 1000 m a.s.l. tem, in shaping the physical characteristics of both lakes and to 1.9 species per lake for lakes at or above 2600 m a.s.l. streams in regard to patterns of loading, transport, storage, However, decoupling the extent that elevation infuences and utilization of inorganic and organic materials (Vannote lake basin characteristics and biogeochemical processes et al. 1980; Kratz et al. 1997; Martin and Soranno 2006). and how these ambient conditions interactively afect the Notably, the Landscape Position Model (LPM) postulates biological properties and community structures, is rela- that in temperate lake systems, downward shifts in land- tively understudied, especially in regards to zooplankton scape position are associated with larger lake size, increased (Vadeboncoeur et al. 2002; Cole et al. 2011). hydrologic connectivity, longer water residence times, and To better understand how the physical and biological increased fsh species richness (Kratz et al. 1997; Martin characteristics that covary with elevation shape alpine lake and Soranno 2006; Soranno et al. 2010). Application of zooplankton communities, we conducted repeat sampling of these and related frameworks have helped capture varia- mountain lakes along an elevation gradient (2480–3550 m tion among mountain lakes or across stream networks with a.s.l.) within Colorado’s Rocky Mountains, USA. Our spe- respect to nutrient concentrations, primary production, and cifc goal was to evaluate the strength and consistency of patterns of biodiversity (Sadro et al. 2012; Epstein et al. correlations between lake elevation and variation in features 2013; Read et al. 2015). For mountain lakes in particular, such as nutrient concentrations, thermal stratifcation and the landscape position of a lake relative to forest treeline has planktonic communities (across multiple trophic levels). enormous potential to afect in-lake ecological processes. Lakes were selected to encompass a range in altitude along- For instance, both the extent and chemical quality of dis- side a representatively broad gradient in lake size, depth, and solved organic matter (DOM) derived from the surrounding catchment area. We used a mixed-modeling framework to landscape sharply infuence the attenuation of UV radiation assess how measured characteristics shifted with elevation (McKnight et al. 1997). Allochthonous DOM derived from while accounting for lake-level characteristics (e.g., deep- terrestrial vegetation and soils is typically enriched in yel- est depth, surface area, and fsh presence) and sources of low-colored, humic materials, which strongly absorb visible autocorrelation (e.g., catchment identity, lake identity and and UV radiation. However, clear water lakes above treeline sampling period). As our survey area encompassed 10 dif- (i.e., alpine) tend to support sparse terrestrial vegetation and ferent catchments and had a meridional extent of 36 km, we thus have lower DOM concentrations often dominated by also compared the relative infuence of geographic distance autochthonous (derived from within-lake processes) rather and elevation on the taxonomic composition of zooplank- than allochthonous inputs (McKnight et al. 1994; Hood ton and phytoplankton among lakes. Building on previous et al. 2003b). As a result, UVR often penetrates deeper in research related to landscape position (1) we expected eleva- the water column, and the carbon subsidy ofered by terres- tion and the associated, patterns of surrounding vegetation, trially derived DOM is typically lacking or much reduced UVR, and ice-free period, to act as a niche flter, leading (e.g., Miller et al. 2009; Sadro et al. 2011). to communities characterized by lower taxonomic rich- Concomitantly, these extreme and dynamic physi- ness and lower population densities. Additionally, (2) we cal characteristics of high-elevation ecosystems likely hypothesized that the diference in elevation between sites exert strong infuences on the biological communities would more strongly relate to community similarity com- of mountain lakes. Compared with low-elevation lakes, pared with straight-line distance or watershed identity alone, alpine lake ecosystems generally support lower species owing to the large number of physical variables that covary richness (Füreder et al. 2006; Fjellheim et al. 2009) and with elevation. The current study ofers a novel contribution have simplifed food webs, often with fewer than three by characterizing how a broad range of variables, including trophic levels (Ward et
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