Projection of invertebrate populations in the headwater streams of a temperate catchment under a changing climate
Graphical abstract
Introduction
The Intergovernmental Panel on Climate Change (IPCC) anticipated that air temperature would increase by ~4 °C in the end of the 21st century (IPCC, 2007), with wide-ranging consequences that include fluctuating precipitation. Changing climatic conditions are expected to cause major stress to stream organisms, through altering flow regimes and elevating water temperature (Sala et al., 2000; Van Vliet et al., 2013). However, climate change-related impacts on stream organisms remain poorly understood (Durance and Ormerod, 2007; Moritz and Agudo, 2013). In addition, the extinction rate in freshwater ecosystems is faster than that in marine and terrestrial ecosystems (Jenkins, 2003; Revenga et al., 2005; Xenopoulos et al., 2005; Strayer and Dudgeon, 2010). Therefore, it is essential to evaluate how future temperature increases due to global warming will affect freshwater ecosystems.
Stream invertebrates are generally poor dispersers that strongly depend on local environments; thus, the relationship between local environments, including water quality and invertebrates, have been frequently studied (e.g., Kerans and Karr, 1994; Glendell et al., 2014). Of the local environmental factors in rivers, water temperature is the primary factor constraining the growth, metabolism, and survival of stream invertebrates (Sweeney and Vannote, 1986; Brittain, 1991; Robinson and Minshall, 1998). Several studies (e.g., Gaufin and Hern, 1971; Quinn et al., 1994) have attempted experimental approaches to understand biological response of invertebrate changes to water temperature, but such research is limited to a few taxonomic groups and cannot inform us about ecological responses at a community level. Researchers (e.g., Statzner and Higler, 1986; Tsuruishi, 2006; Duggan et al., 2007) have explored thermal influences on invertebrates through field monitoring; however, habitats of stream invertebrates are determined by complex environmental factors, including flow velocity, substrate, and water quality. Consequently, it is difficult to detect the specific impact of water temperature on the invertebrate community of streams.
In general, field studies of invertebrate in this line of inquiry have focused on midstream to downstream zones because of the relative ease of entry to a watercourse for sampling. However, midstream to downstream water quality is generally impaired by point and nonpoint source pollution, which might obscure the link between water temperature and invertebrates. In the upland forests, stream water is mainly composed of intermediate flow filtered through forested soil (Neary et al., 2009), with consumption and canopy interception reducing direct runoff (Hewlett, 1982). Typically, water quality is higher in the headwaters of such rivers than downstream, while water temperature varies depending on the altitude and season in temperate catchments (Sponseller et al., 2001). Furthermore, headwater streams are generally free of thermal pollution caused by industrial effluents (Caissie, 2006). Such environments are therefore suitable study areas for improving our understanding of ecological responses to water temperature gradients (Arai et al., 2015).
To date, researchers have modeled the impact of climatic-change on stream invertebrates using general circulation models (GCMs). Li et al. (2013) used invertebrate data collected throughout South Korea and predicted the habitat distribution of invertebrates under future climates by introducing GCM-projected air temperatures. However, this study assumed negligible effects from other habitat conditions (e.g., current velocity). In addition to air temperature, other reports have used precipitation and topographic variables to project the distribution of stream invertebrates under changing climate scenarios (Bálint et al., 2011; Domisch et al., 2011; Domisch et al., 2013). Although previous studies succeeded in integrating GCMs into habitat models, challenges remain. First, surrogates of water temperature measures (e.g., annual maximum air temperature) were used because air temperature variables are easily accessible. Li et al., 2013, Li et al., 2014 projected habitat distribution of stream invertebrates by water temperature although the water temperature used was estimated by GCM-projected air temperature using a linear regression model. Second, projection uncertainty under different GCMs (Wilby and Harris, 2006) is rarely taken into account, with only one or two GCMs generally being used. This uncertainty could lead to bias when attempting to project stream invertebrate responses. Thus, the number of GCMs should be increased for a more objective evaluation of means and deviations.
We aimed to develop a predictive model of stream invertebrate density under different climatic change scenarios using multiple GCMs, biological field data from headwater streams, and a hydrological simulation (see conceptual study flow in Fig. 1). We used existing quantitative sampling data of stream invertebrate and water temperature (Arai et al., 2015) in the headwater streams of a temperate catchment in northeast Japan. The headwater streams selected for the current study had similar hydraulic and geomorphic conditions and was free of human-induced impacts. In addition, stream invertebrates are well known to respond to environmental changes. Thus, these data are useful for projecting fate of headwater invertebrates in response to the water temperature rise while minimizing the effects of driving factors other than water temperature on invertebrates. We adopted eight GCM types and three RCP (Representative Concentration Pathways) scenarios to check model uncertainty and assess climate-change effects on stream invertebrate density. By inputting GCM-projected air temperature into an existing distributed hydrological-thermal model (hereafter hydrothermal model), we computed stream water temperatures for the studied headwaters. Finally, regression models were developed to project invertebrate density at the study catchment under different climate change scenarios (based on water temperature). Our results are expected to identify invertebrate groups appropriate for use as thermal-sensitive indicators of climate change, which could then be used in the predictions of climate-change effects over much broader scales.
Section snippets
Study area
We studied headwater streams in the Natori River catchment (939 km2), northeast Japan. The annual mean air temperature and annual precipitation were 9.8 °C and 1593.5 mm at the Nikkawa Meteorological Station (265 m above sea level [a.s.l.]), 12.7 °C and 1179.5 mm at the Sendai Meteorological Station (39 m a.s.l.), and 11.2 °C and 1257.5 mm at the Zao Meteorological Station (112 m a.s.l.), respectively (Fig. 2). Surface air temperature and precipitation seasonally varies (Fig. S1). Gunawardhana
Results
Increases in projected air temperature were significantly greater under higher radiative forcing scenarios in both future periods (paired t-test, P < 0.001), with the exception of no significant difference being observed between RCP2.6 and RCP4.5 in the near future. In addition, temperature increases under the RCP4.5 and RCP8.5 scenarios in the far future were significantly higher than those in the near future (paired t-test, P < 0.001).
Seven out of the eight higher taxonomic groups exhibited
Discussion
The present study combined a physically-based hydrothermal model, GCMs, and regressions to predict the density of invertebrate based on water temperature in headwater streams. We successfully projected the population density of stream invertebrates under different climate-change scenarios. Previous studies faced various challenges in predicting the distribution/population density of stream animals, including invertebrates. For instance, surrogate variables of water temperature (e.g., air
Conclusions
The present study integrated a physically based hydrological-thermal simulation, eight general circulation models, and headwater invertebrate samples to project invertebrate population density under different climate change scenarios. In brief, our study successfully projected headwater temperature using a hydrothermal simulation. This model was used to project invertebrate densities, select sensitive indicator taxa of climate change, and obtain spatial predictions of declines in the population
Acknowledgements
This research was partially supported by the Ministry of Education, Science, Sports and Culture through a Grant-in-Aid for Scientific Research [grant numbers 16H02363, 16H05750]. We thank the anonymous reviewers for providing constructive comments on our manuscript.
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