Specialized Meltwater Biodiversity Persists Despite Widespread Deglaciation

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Specialized Meltwater Biodiversity Persists Despite Widespread Deglaciation Specialized meltwater biodiversity persists despite widespread deglaciation Clint C. Muhlfelda,b,1,2, Timothy J. Clinea,2, J. Joseph Gierscha,2, Erich Peitzscha, Caitlyn Florentinea, Dean Jacobsenc, and Scott Hotalingd aNorthern Rocky Mountain Science Center, US Geological Survey, Glacier National Park, West Glacier, MT 59936; bFlathead Lake Biological Station, University of Montana, Polson, MT 59860; cFreshwater Biological Section, Department of Biology, University of Copenhagen, 2100 Copenhagen, Denmark; and dSchool of Biological Sciences, Washington State University, Pullman, WA 99164 Edited by Warwick F. Vincent, Laval University, Quebec City, Canada, and accepted by Editorial Board Member David W. Schindler March 30, 2020 (received for review January 29, 2020) Glaciers are important drivers of environmental heterogeneity and We conducted this test in Glacier National Park (GNP), biological diversity across mountain landscapes. Worldwide, gla- Montana, a protected landscape (e.g., World Heritage Site and ciers are receding rapidly due to climate change, with important Biosphere Reserve) that has experienced widespread glacier loss consequences for biodiversity in mountain ecosystems. However, since the end of the Little Ice Age (LIA, ca. 1850; Fig. 1). Like the effects of glacier loss on biodiversity have never been many mountain ranges worldwide, this region harbors an ex- quantified across a mountainous region, primarily due to a lack tensive network of high-elevation streams that are highly heter- of adequate data at large spatial and temporal scales. Here, we ogenous in terms of hydrologic source contributions, temperature, combine high-resolution biological and glacier change (ca. < 1850–2015) datasets for Glacier National Park, USA, to test the and flow regimes across small spatial extents ( 1 km). These habitat prediction that glacier retreat reduces biodiversity in mountain mosaics support diverse biological communities with high levels of ecosystems through the loss of uniquely adapted meltwater endemism, including two stoneflies (Lednia tumana and Zapada stream species. We identified a specialized cold-water invertebrate glacier) that were recently listed under the US Endangered Species community restricted to the highest elevation streams primarily Act exclusively due to climate change-induced glacier and snow loss below glaciers, but also snowfields and groundwater springs. (17, 18). Of the 146 glaciers present at the end of the LIA, only 35% We show that this community and endemic species have unex- persisted through 2005 (19), with complete deglaciation predicted ECOLOGY pectedly persisted in cold, high-elevation sites, even in catchments by 2100 (5). We focused on alpine stream invertebrates because ∼ that have not been glaciated in 170 y. Future projections suggest they are constrained to the highest elevations of aquatic habitats substantial declines in suitable habitat, but not necessarily loss of and are considered a “canary in the coal mine” for climate-induced this community with the complete disappearance of glaciers. Our biodiversity loss in mountain ecosystems (20). They are also key findings demonstrate that high-elevation streams fed by snow and other cold-water sources continue to serve as critical climate components of freshwater communities, performing vital roles in refugia for mountain biodiversity even after glaciers disappear. Significance mountain streams | glacier loss | biodiversity | invertebrate communities | climate change Glaciers are retreating rapidly due to climate change, and these changes are predicted to reduce biodiversity in mountain eco- ountain regions are rich in biodiversity, harboring high systems through the loss of specialized meltwater species. Mlevels of species richness and endemism (1). The genera- However, direct observations of how glacier change affects tion and maintenance of mountain biodiversity ultimately de- biological communities living in mountain streams are scarce. pends on geological and climatic processes that promote habitat Here, we show that a specialized cold-water invertebrate community has unexpectedly persisted in cold, high-elevation heterogeneity across these dynamic landscapes (2). Glaciers and streams despite widespread glacier loss and habitat reductions other meltwater sources enhance habitat heterogeneity of in Glacier National Park since the Little Ice Age (ca. 1850). Al- mountainous regions by shaping local geomorphology and pro- though shrinking habitat and increased fragmentation pose – viding cold meltwater to downstream ecosystems (2 4). How- significant risks to these range-restricted species, our results ever, glaciers are rapidly shrinking and disappearing due to demonstrate that specialized meltwater communities may be global warming (5–8), and these trends are predicted to accel- more resilient to glacier recession than previously thought. erate over the coming decades (5, 9). Glacier loss is considered a major threat to biodiversity in mountain regions worldwide (2, Author contributions: C.C.M., T.J.C., J.J.G., and S.H. designed research; C.C.M., T.J.C., J.J.G., E.P., C.F., D.J., and S.H. performed research; C.C.M., T.J.C., and J.J.G. analyzed data; and 10, 11), yet direct, quantitative tests of how glacier retreat affects C.C.M., T.J.C., J.J.G., E.P., C.F., D.J., and S.H. wrote the paper. biodiversity across a range of spatial scales are scarce (12, 13). The authors declare no competing interest. The rapid decline of mountain glaciers reduces meltwater This article is a PNAS Direct Submission. W.F.V. is a guest editor invited by the contributions to downstream ecosystems, altering flow regimes, Editorial Board. increasing water temperatures, and ultimately homogenizing This open access article is distributed under Creative Commons Attribution-NonCommercial- stream habitats (3, 14). Glacier loss is expected to increase local NoDerivatives License 4.0 (CC BY-NC-ND). diversity within streams as more diverse, warmer-water com- Data deposition: The stream invertebrate data and contemporary glacier data used in this munities shift upstream, but decrease regional diversity as study are available online from the USGS in ScienceBase (https://doi.org/10.5066/ P9RCMMKL). The R code for all statistical analyses used in this study have been deposited unique glacier-dependent species and communities are lost (15, in GitHub (https://github.com/tjcline/AlpineBugs_PNAS2020). – 16). Global predictions suggest that 11 38% of species will be 1To whom correspondence may be addressed. Email: [email protected]. lost following complete disappearance of glaciers in a region 2C.C.M., T.J.C., and J.J.G. contributed equally to this work. (15). Here, we explicitly test the prediction that glacier recession This article contains supporting information online at https://www.pnas.org/lookup/suppl/ reduces stream biodiversity through the loss of specialized doi:10.1073/pnas.2001697117/-/DCSupplemental. meltwater species across a large mountainous region. www.pnas.org/cgi/doi/10.1073/pnas.2001697117 PNAS Latest Articles | 1of7 Downloaded by guest on September 28, 2021 Fig. 1. Glaciers have rapidly receded in GNP since the LIA. (A) Map and histogram of glacial loss for all 129 sampling sites (colored circles) since the end of the LIA (ca. 1850). White to dark shading represents topographic relief. Sampling sites (red points), LIA glacier extent (yellow), and historical photos of glacier loss in Sperry (B), Boulder (C), Grinnell (D) basins are shown. Sites where glaciers were absent in LIA are not included in the histogram. ecosystem functioning, such as organic matter decomposition and reduced glacial cover and warmer water temperatures (Fig. 2). nutrient cycling. These findings are consistent with previous studies (15, 16, 24, 25). Using high-resolution satellite imagery, we quantified change Taxonomic differences (dissimilarity) between individual sites and in glacier cover by mapping the area of present-day glaciers (ca. the regional species pool (β-diversity) was higher among sites with 2015) and analyzing their maximum at the end of the LIA (18, greater glacial cover and colder water temperatures. Spatial turn- 19, 21). We quantified the abundance and diversity of stream over of taxa accounted for 98.5% of overall beta-diversity among invertebrate communities at 129 sites (within and across streams) sites, demonstrating that glaciated reaches support distinct com- in 36 catchments with varying levels of contemporary glacial munities (SI Appendix,TableS2). These results also indicate that influence (percentage of present-day glacier cover, 0–91%) and local increases in species diversity may come at the expense of re- glacier loss since the LIA (8–100%) (22). Sample sites spanned a gional species diversity as glacier-dependent species are lost and range of water sources (glacier, snowfield, and groundwater high-elevation streams become environmentally and taxonomically spring), summer water temperatures (1–10 °C), elevations (1,507– homogenized (15). However, there was significant variation in com- 2,431 m), and distances to source (1–3,853 m). We identified dis- munity diversity among sites without glaciers, suggesting environ- tinct invertebrate communities using latent Dirichlet allocation mental conditions supporting distinct communities may exist (LDA) (23), a probabilistic hierarchical cluster analysis, with no a elsewhere on the landscape. priori habitat information included. We then evaluated how a suite We identified a total of 113 taxa
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