Controls on ecosystem water-use and water-use efficiency: Insights from a comparison between grassland and riparian forest in the northern Great Plains
Introduction
The Great Plains is the second largest biome in North America and extends south from the aspen parkland and boreal forest areas in Alberta and Saskatchewan, Canada through the USA to New Mexico and Texas (Ostlie et al., 1997; Zhang et al., 2011). There is a substantial west to east gradient in precipitation that divides the native vegetation of the Great Plains into short-, mid- and tall grasslands or prairie ecosystems (Heisler-White et al., 2009; Wilcox et al., 2015). Productivity and water-use in prairie grassland ecosystems are strongly controlled by the amount of summer precipitation inputs, although the size of rain events and the interval between rain events can also significantly influence grassland ecosystem physiology (Knapp and Smith, 2001; Heisler-White et al., 2008, 2009; Knapp et al., 2008). Within this grassland-dominated landscape, riparian ecosystems also exist along the floodplains of rivers that distribute snow-melt water from the Rocky Mountains as they flow out through the prairies (Rood et al., 2003). Within the northern Great Plains of southern Alberta, river water also flows laterally out of the main river channels (“losing rivers”) creating an alluvial aquifer below the river floodplain (Hauer et al., 2016). A variety of important ecological processes connect the aquatic and terrestrial ecosystems through this alluvial aquifer and these ecological interactions promote regional biodiversity across the full spectrum from microbes to vertebrate animals (Hauer et al., 2016). Riparian cottonwood forests develop on these floodplains within the semiarid landscape that is otherwise treeless, in part supported by the alluvial groundwater which supplements any precipitation inputs to the soil (Snyder and Williams, 2000; Rood et al., 2003; Scott et al., 2003, 2000; Flanagan et al., 2017). The riparian forest ecosystems contribute many valued ecosystem services (Naiman et al., 2005; Hauer et al., 2016), but their health and survival has been threatened by reductions in river flows associated with dams that divert river water for agricultural irrigation, municipal and industrial consumption (Rood et al., 1995, 2003, 2005, 2008; Schindler and Donahue, 2006). This raises the fundamental question, how much river water is required to support healthy riparian forest ecosystems within the Great Plains biome? In order to help answer this question, and to provide some ecological perspective on the water-use requirements of riparian cottonwood forest ecosystems, our major objective in this study was to compare water-use and water-use efficiency between a native grassland ecosystem and a native, riparian cottonwood ecosystem in the northern Great Plains of southern Alberta. This comparison should provide insights into the amount of river water required to support a healthy riparian forest ecosystem in this region, as the local grasslands were expected to rely only on water sourced from summer precipitation inputs (Wever et al., 2002). More generally, this analysis would allow evaluation of some important factors influencing water-use and water-use efficiency in these two ecosystems.
In this paper we define water-use as cumulative ecosystem evapotranspiration (ET) during the growing season, and water-use efficiency (WUE) as the ratio of ecosystem photosynthesis to ecosystem ET. Several ecological and physiological factors can control ecosystem water-use and WUE, including: leaf area index (LAI), stomatal conductance, precipitation input, soil moisture content, access to groundwater, temperature, vapor pressure difference (VPD), and growing season length (Kelliher et al., 1995; Ponton et al., 2006; Beer et al., 2009). We predicted that cottonwood riparian forests and native grassland ecosystems should differ for several of these factors, as is described below. The contrasting dominant plant functional types associated with cottonwood forests and native prairie grassland will result in important differences in leaf and canopy photosynthetic gas exchange processes. Broad-leaf deciduous trees tend to have lower ratios of stomatal conductance to photosynthetic capacity than C3 grasses (Smedley et al., 1991; Brooks et al., 1997), but the typically larger LAI of forests than grasslands results in canopy conductance being higher in the forest ecosystems (Kelliher et al., 1995; Wever et al., 2002; Flanagan et al., 2017). While precipitation input will be the same in nearby riparian forest and grassland ecosystems, groundwater access by deep-rooted, phreatophytic cottonwood trees should increase the supply of water to riparian forests above that available to grasslands (Scott et al., 2003, 2004). In addition, the depth of soil and permeable substrate beneath floodplain forests (2–3 m) exceeds that of native grasslands in the northern Great Plains (0–30 cm), and so the riparian forests should have greater soil water storage capacity. Local forest and grassland ecosystems are exposed to similar aerial environmental conditions (photosynthetically active radiation, temperature, VPD), but they differ in ecological strategies of the dominant plant functional types. In addition, the differing amounts of available soil water can result in contrasting growing season lengths for forest and grassland ecosystems. Grassland plants grow fast and use water quickly when it is readily available and then go dormant to survive times of water shortages (Flanagan and Adkinson, 2011). This ecological strategy of grassland plants can result in significantly shorter active growing seasons than is apparent for riparian forest ecosystems.
In this study, we compared ecosystem water-use and WUE between nearby riparian cottonwood forest and native grassland ecosystems in southern Alberta using eddy covariance measurements. Our goal was to test these predictions made above about mechanistic controls on ecosystem water-use, and to provide insights into the amount of alluvial groundwater and stored soil water that is required to support a healthy riparian forest ecosystem within the Great Plains biome. The study was conducted during May-September in three different years that had contrasting precipitation, air temperature and VPD, and so provided perspective on summertime environmental controls that influence water-use and WUE in riparian forest and grassland ecosystems in the northern Great Plains biome.
Section snippets
Study site descriptions
The two study sites were located in Lethbridge, Alberta, Canada near the northwestern limit of the Great Plains biome. The mean annual temperature for Lethbridge was 5.7 °C and average annual precipitation was 386.3 mm (1971-2010: Canadian Climate Normals, Environment Canada; climate.weather.gc.ca/climate_normals/). The forest site was a riparian cottonwood forest in the Helen Schuler Nature Reserve (HSNR, 49.702 °N, 112.863 °W, elevation 928 m) within the Oldman River valley (Flanagan et al.,
Comparison of environmental conditions
Total growing season (May-September) precipitation was higher in 2014 (363 mm) than normal (258 ± 106 mm, 30-year average ± SD), near normal in 2015 (181 mm), and below normal in 2017 (125 mm) (Fig. 1). Extremely high rainfall of 144.5 mm during days 164–170 caused widespread over-bank flooding of the Oldman River in 2014 (Fig. 2). This flood was typical for floods of the area in that it was caused by heavy precipitation during late May to early July when soils of the upper watershed catchment
Discussion
This comparative study between native grassland and riparian forest ecosystems, which were exposed to the same aboveground environmental conditions, allowed for evaluation of other dominant factors including plant functional type, LAI and soil and groundwater access for their effects on ecosystem water-use and WUE. In addition, the contrasting environmental conditions among study years provided insights into mechanisms that control water-use and WUE in the two ecosystems. Such comparative
Acknowledgements
This research was part of the Functional Flows: A Practical Strategy for Healthy Rivers project and was supported by grants from Alberta Innovates, the Natural Sciences and Engineering Council of Canada - Discovery Grant Program, and Conoco Phillips Canada. We thank Rachel Tkach, Lauren Schlerloski, Dylan Nikkel, Eric Sharp, Emily Wilton, Kayla Johnson, Caitlin Pelletier, Tyler Tremel, and David Pearce for help with some of the field and lab work. David Ellis (City of Lethbridge) and Coreen
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