1. Hock, R. et al. in IPCC Special Report on the Ocean and Cryosphere in a Changing Climate (eds Pörtner, H.-O. et al.) Ch. 2 (IPCC, 2019).

2. Zemp, M. et al. Global glacier mass changes and their contributions to sea-level rise from 1961 to 2016. Nature 568, 382–386 (2019).

3. Sorg, A., Bolch, T., Stoffel, M., Solomina, O. & Beniston, M. Climate change impacts on glaciers and runoff in the Tien Shan (Central Asia). Nat. Clim. Change 2, 725–731 (2012).

4. Milner, A. M. et al. Glacier shrinkage driving global changes in downstream systems. Proc. Natl Acad. Sci. USA 114, 9770–9778 (2017).

5. Lee, J. R. et al. Climate change drives expansion of Antarctic ice-free habitat. Nature 547, 49–54 (2017).

6. Jacobsen, D., Milner, A. M., Brown, L. E. & Dangles, O. Biodiversity under threat in glacier-fed river systems. Nat. Clim. Change 2, 361–364 (2012).

7. Rabatel, A. et al. Current state of glaciers in the tropical Andes: a multi-century perspective on glacier evolution and climate change. Cryosphere 7, 81–102 (2013).

8. Beniston, M. et al. The European mountain cryosphere: a review of its current state, trends, and future challenges. Cryosphere 12, 759–794 (2018).

9. Huss, M. et al. Toward mountains without permanent snow and ice. Earth’s Future 5, 418–435 (2017).

10. Bell, E. M. Life at Extremes: Environments, Organisms and Strategies for Survival (CABI, 2012).

11. Grange, L. J. & Smith, C. R. Megafaunal communities in rapidly warming fjords along the West Antarctic Peninsula: hotspots of abundance and beta diversity. PLoS ONE 8, e77917 (2013).

12. Cauvy-Fraunié, S., Espinosa, R., Andino, P., Jacobsen, D. & Dangles, O. Invertebrate metacommunity structure and dynamics in an Andean glacial stream network facing climate change. PLoS ONE 10, e0136793 (2015).

13. Munoz, N. J., Farrell, A. P., Heath, J. W. & Neff, B. D. Adaptive potential of a Pacific salmon challenged by climate change. Nat. Clim. Change 5, 163–166 (2015).

14. Milner, A. M., Taylor, R. C. & Winterbourn, M. J. Longitudinal distribution of macroinvertebrates in two glacier‐fed New Zealand rivers. Freshw. Biol. 46, 1765–1775 (2001).

15. Ronowicz, M., Włodarska-Kowalczuk, M. & Kukliński, P. Patterns of hydroid (Cnidaria, Hydrozoa) species richness and distribution in an Arctic glaciated fjord. Polar Biol. 34, 1437–1445 (2011).

16. Gobbi, M., Isaia, M. & De Bernardi, F. Arthropod colonisation of a debris-covered glacier. Holocene 21, 343–349 (2011).

17. Giersch, J. J., Hotaling, S., Kovach, R. P., Jones, L. A. & Muhlfeld, C. C. Climate‐induced glacier and snow loss imperils alpine stream insects. Glob. Change Biol. 23, 2577–2589 (2017).

18. Jacobsen, D. & Dangles, O. Environmental harshness and global richness patterns in glacier‐fed streams. Glob. Ecol. Biogeogr. 21, 647–656 (2012).

19. Milner, A. M., Fastie, C. L., Chapin, F. S., Engstrom, D. R. & Sharman, L. C. Interactions and linkages among ecosystems during landscape evolution. BioScience 57, 237–247 (2007).

20. Zawierucha, K., Kolicka, M., Takeuchi, N. & Kaczmarek, Ł. What animals can live in cryoconite holes? A faunal review. J. Zool. 295, 159–169 (2015).

21. Cauvy-Fraunié, S. et al. Ecological responses to experimental glacier-runoff reduction in alpine rivers. Nat. Commun. 7, 12025 (2016).

22. Arimitsu, M. L. et al. Distribution and spawning dynamics of capelin (Mallotus villosus) in Glacier Bay, Alaska: a cold water refugium. Fish. Oceanogr. 17, 137–146 (2008).

23. Uehlinger, U., Robinson, C., Hieber, M. & Zah, R. The physico-chemical habitat template for periphyton in alpine glacial streams under a changing climate. Hydrobiologia 657, 107–121 (2010).

24. Dolezal, J. et al. Primary succession following deglaciation at Koryto Glacier Valley, Kamchatka. Arct. Antarct. Alp. Res. 40, 309–322 (2008).

25. Weslawski, J. & Legezynska, J. Glaciers caused zooplankton mortality? J. Plankton Res. 20, 1233–1240 (1998).

26. Řeháková, K., Stibal, M., Šabacká, M. & Řehák, J. Survival and colonisation potential of photoautotrophic microorganisms within a glacierised catchment on Svalbard, High Arctic. Polar Biol. 33, 737–745 (2010).

27. Wilhelm, L., Singer, G. A., Fasching, C., Battin, T. J. & Besemer, K. Microbial biodiversity in glacier-fed streams. ISME J. 7, 1651–1660 (2013).

28. Park, S. J. et al. Influence of deglaciation on microbial communities in marine sediments off the coast of Svalbard, Arctic Circle. Microb. Ecol. 62, 537–548 (2011).

29. Breen, K. & Levesque, E. Proglacial succession of biological soil crusts and vascular plants: biotic interactions in the High Arctic. Can. J. Bot. 84, 1714–1731 (2006).

30. Oehl, F., Schneider, D., Sieverding, E. & Burga, C. A. Succession of arbuscular mycorrhizal communities in the foreland of the retreating Morteratsch Glacier in the Central Alps. Pedobiologia 54, 321–331 (2011).

31. Malard, F., Lafont, M., Burgherr, P. & Ward, J. A comparison of longitudinal patterns in hyporheic and benthic oligochaete assemblages in a glacial river. Arct. Antarct. Alp. Res. 33, 457–466 (2001).

32. Schlegel, J. & Riesen, M. Environmental gradients and succession patterns of carabid beetles (Coleoptera: Carabidae) in an Alpine glacier retreat zone. J. Insect Conserv. 16, 657–675 (2012).

33. Seimon, T. A. et al. Upward range extension of Andean anurans and chytridiomycosis to extreme elevations in response to tropical deglaciation. Glob. Change Biol. 13, 288–299 (2007).

34. Arendt, K. E., Nielsen, T. G., Rysgaard, S. & Tönnesson, K. Differences in plankton community structure along the Godthåbsfjord, from the Greenland Ice Sheet to offshore waters. Mar. Ecol. Prog. Ser. 401, 49–62 (2010).

35. Berge, J. et al. Changes in the decapod fauna of an Arctic fjord during the last 100 years (1908–2007). Polar Biol. 32, 953–961 (2009).

36. Błażewicz-Paszkowycz, M. & Sekulska-Nalewajko, J. Tanaidacea (Crustacea, Malacostraca) of two polar fjords: Kongsfjorden (Arctic) and Admiralty Bay (Antarctic). Polar Biol. 27, 222–230 (2004).

37. Carney, D., Oliver, J. S. & Armstrong, C. Sedimentation and composition of wall communities in Alaskan fjords. Polar Biol. 22, 38–49 (1999).

38. Day, R. H. & Nigro, D. A. Feeding ecology of Kittlitz’s and marbled murrelets in Prince William Sound, Alaska. J. Waterbirds Soc. 23, 1–14 (2000).

39. De Skowronski, R. S. & Corbisier, T. N. Meiofauna distribution in Martel Inlet, King George Island (Antarctica): sediment features versus food availability. Polar Biol. 25, 126–134 (2002).

40. De Skowronski, R. S. et al. Distribution of microphytobenthic biomass in Martel Inlet, King George Island (Antarctica). Polar Biol. 32, 839–851 (2009).

41. Etherington, L. L., Hooge, P. N., Hooge, E. R. & Hill, D. F. Oceanography of Glacier Bay, Alaska: implications for biological patterns in a glacial fjord estuary. Estuar. Coasts 30, 927–944 (2007).

42. Fetzer, I., Lønne, O. & Pearson, T. The distribution of juvenile benthic invertebrates in an arctic glacial fjord. Polar Biol. 25, 303–315 (2002).

43. Gontar, V. I., Hop, H. & Voronkov, A. Y. Diversity and distribution of Bryozoa in Kongsfjorden, Svalbard. Pol. Polar Res. 22, 187–204 (2001).

44. Grzelak, K. & Kotwicki, L. Meiofaunal distribution in Hornsund fjord, Spitsbergen. Polar Biol. 35, 269–280 (2012).

45. Hald, M. & Korsun, S. Distribution of modern benthic foraminifera from fjords of Svalhard, European Arctic. J. Foramin. Res. 27, 101–122 (1997).

46. Jankowska, K. & Wieczorek, P. Abundance and biomass of bacteria in two Arctic glacial fjords. Pol. Polar Res. 26, 77–84 (2005).

47. Kaczmarek, H., Włodarska-Kowalczuk, M., Legezynska, J. & Zajaczkowski, M. Shallow sublittoral macrozoobenthos in Kongsfjord, west Spitsbergen, Svalbard. Pol. Polar Res. 26, 137–155 (2005).

48. Keck, A., Wiktor, J., Hapter, R. & Nilsen, R. Phytoplankton assemblages related to physical gradients in an arctic, glacier-fed fjord in summer. ICES J. Mar. Sci. 56, 203–214 (1999).

49. Kędra, M., Włodarska-Kowalczuk, M. & Węsławski, J. M. Decadal change in macrobenthic soft-bottom community structure in a high Arctic fjord (Kongsfjorden, Svalbard). Polar Biol. 33, 1 (2010).

50. Kędra, M., Legeżyńska, J. & Walkusz, W. Shallow winter and summer macrofauna in a high Arctic fjord (79 N, Spitsbergen). Mar. Biodiv. 41, 425–439 (2011).

51. Kędra, M., Pabis, K., Gromisz, S. & Węsławski, J. M. Distribution patterns of polychaete fauna in an Arctic fjord (Hornsund, Spitsbergen). Polar Biol. 36, 1463–1472 (2013).

52. Kissling, M. L., Reid, M., Lukacs, P. M., Gende, S. M. & Lewis, S. B. Understanding abundance patterns of a declining seabird: implications for monitoring. Ecol. Appl. 17, 2164–2174 (2007).

53. Korsun, S., Pogodina, I., Forman, S. & Lubinski, D. Recent foraminifera in glaciomarine sediments from three Arctic fjords of Novaja Zemlja and Svalbard. Polar Res. 14, 15–32 (1995).

54. Korsun, S. & Hald, M. Modern benthic foraminifera off Novaya Zemlya tidewater glaciers, Russian Arctic. Arct. Alp. Res. 30, 61–77 (1998).

55. Korsun, S. & Hald, M. Seasonal dynamics of benthic foraminifera in a glacially fed fjord of Svalbard, European Arctic. J. Foramin. Res. 30, 251–271 (2000).

56. Kotwicki, L., Szymelfenig, M., De Troch, M. & Zajaczkowski, M. Distribution of meiofauna in Kongsfjorden, Spitsbergen. Polar Biol. 27, 661–669 (2004).

57. Kuklinski, P. Fauna of Bryozoa from Kongsfjorden, West Spitsbergen. Pol. Polar Res. 23, 193–206 (2002).

58. Kuklinski, P., Gulliksen, B., Lønne, O. J. & Weslawski, J. M. Composition of bryozoan assemblages related to depth in Svalbard fjords and sounds. Polar Biol. 28, 619–630 (2005).

59. Kuklinski, P., Gulliksen, B., Lønne, O. J. & Weslawski, J. M. Substratum as a structuring influence on assemblages of Arctic bryozoans. Polar Biol. 29, 652–661 (2006).

60. Kwasniewski, S., Hop, H., Falk-Petersen, S. & Pedersen, G. Distribution of Calanus species in Kongsfjorden, a glacial fjord in Svalbard. J. Plankton Res. 25, 1–20 (2003).

61. Legeżyńska, J. et al. The malacostracan fauna of two Arctic fjords (west Spitsbergen): the diversity and distribution patterns of its pelagic and benthic components. Oceanologia 59, 541–564 (2017).

62. Majewski, W. Benthic foraminiferal communities: distribution and ecology in Admiralty Bay, King George Island, West Antarctica. Pol. Polar Res. 26, 159–214 (2005).

63. Majewski, W. & Olempska, E. Recent ostracods from Admiralty Bay, King George Island, West Antarctica. Pol. Polar Res. 26, 13–36 (2005).

64. Majewski, W., Pawlowski, J. & Zajaczkowski, M. Monothalamous foraminifera from West Spitsbergen fjords, Svalbard: a brief overview. Pol. Polar Res. 26, 269–285 (2005).

65. Moon, H.-W., Hussin, W. M. R. W., Kim, H.-C. & Ahn, I.-Y. The impacts of climate change on Antarctic nearshore mega-epifaunal benthic assemblages in a glacial fjord on King George Island: responses and implications. Ecol. Ind. 57, 280–292 (2015).

66. Murray, C. et al. The influence of glacial melt water on bio-optical properties in two contrasting Greenlandic fjords. Estuar. Coast. Shelf Sci. 163, 72–83 (2015).

67. Mutschke, E. & Gorny, M. The benthic decapod fauna in the channels and fjords along the South Patagonian Icefield, Southern Chile. Sci. Mar. 63, 315–319 (1999).

68. Okolodkov, Y. B., Hapter, R. & Semovski, S. V. Phytoplankton in Kongsfjorden, Spitsbergen, July 1996. Sarsia 85, 345–352 (2000).

69. Pabis, K. & Sicinski, J. Distribution and diversity of polychaetes collected by trawling in Admiralty Bay: an Antarctic glacial fiord. Polar Biol. 33, 141–151 (2010).

70. Pabis, K., Sicinski, J. & Krymarys, M. Distribution patterns in the biomass of macrozoobenthic communities in Admiralty Bay (King George Island, South Shetlands, Antarctic). Polar Biol. 34, 489–500 (2011).

71. Pabis, K., Hara, U., Presler, P. & Sicinski, J. Structure of bryozoan communities in an Antarctic glacial fjord (Admiralty Bay, South Shetlands). Polar Biol. 37, 737–751 (2014).

72. Pabis, K., Kędra, M. & Gromisz, S. Distinct or similar? Soft bottom polychaete diversity in Arctic and Antarctic glacial fjords. Hydrobiologia 742, 279–294 (2015).

73. Pabis, K. & Sobczyk, R. Small-scale spatial variation of soft-bottom polychaete biomass in an Antarctic glacial fjord (Ezcurra Inlet, South Shetlands): comparison of sites at different levels of disturbance. Helgol. Mar. Res. 69, 113–121 (2015).

74. Pugh, P. J. A. & Davenport, J. Colonisation vs. disturbance: the effects of sustained ice-scouring on intertidal communities. J. Exp. Mar. Biol. Ecol. 210, 1–21 (1997).

75. Renaud, P. E. et al. Multidecadal stability of benthic community structure in a high-Arctic glacial fjord (van Mijenfjord, Spitsbergen). Polar Biol. 30, 295–305 (2007).

76. Ronowicz, M. Species diversity of Arctic gravel beach: case study for species poor habitats. Pol. Polar Res. 26, 287–297 (2005).

77. Ronowicz, M., Włodarska-Kowalczuk, M. & Kuklinski, P. Factors influencing hydroids (Cnidaria: Hydrozoa) biodiversity and distribution in Arctic kelp forest. J. Mar. Biol. Assoc. UK 88, 1567–1575 (2008).

78. Sabbatini, A., Morigi, C., Negri, A. & Gooday, A. J. Distribution and biodiversity of stained monothalamous foraminifera from Tempelfjord, Svalbard. J. Foramin. Res. 37, 93–106 (2007).

79. Sejr, M. K., Włodarska-Kowalczuk, M., Legeżyńska, J. & Blicher, M. E. Macrobenthic species composition and diversity in the Godthaabsfjord system, SW Greenland. Polar Biol. 33, 421–431 (2010).

80. Siciński, J., Pabis, K., Jażdżewski, K., Konopacka, A. & Błażewicz-Paszkowycz, M. Macrozoobenthos of two Antarctic glacial coves: a comparison with non-disturbed bottom areas. Polar Biol. 35, 355–367 (2012).

81. Taggart, S., Hooge, P., Mondragon, J., Hooge, E. & Andrews, A. Living on the edge: distribution of Dungeness crab Cancer magister in a recently deglaciated fjord. Mar. Ecol. Prog. Ser. 246, 241–252 (2003).

82. Tikhonenkov, D. Species diversity and changes of communities of heterotrophic flagellates (protista) in response to glacial melt in King George Island, the South Shetland Islands, Antarctica. Antarct. Sci. 26, 133–144 (2014).

83. Urban-Malinga, B., Wiktor, J., Jabłońska, A. & Moens, T. Intertidal meiofauna of a high-latitude glacial Arctic fjord (Kongsfjorden, Svalbard) with emphasis on the structure of free-living nematode communities. Polar Biol. 28, 940–950 (2005).

84. Voronkov, A., Stepanjants, S. D. & Hop, H. Hydrozoan diversity on hard bottom in Kongsfjorden, Svalbard. J. Mar. Biol. Assoc. UK 90, 1337–1352 (2010).

85. Voronkov, A. & Hop, H. & Gulliksen, B. Diversity of hard-bottom fauna relative to environmental gradients in Kongsfjorden, Svalbard. Polar Res. 32, 11208 (2013).

86. Walkusz, W. et al. Seasonal and spatial changes in the zooplankton community of Kongsfjorden, Svalbard. Polar Res. 28, 254–281 (2009).

87. Wang, G., Guo, C., Luo, W., Cai, M. & He, J. The distribution of picoplankton and nanoplankton in Kongsfjorden, Svalbard during late summer 2006. Polar Biol. 32, 1233–1238 (2009).

88. Weslawski, J. M., Wiktor, J. & Kotwicki, L. Increase in biodiversity in the arctic rocky littoral, Sorkappland, Svalbard, after 20 years of climate warming. Mar. Biodiv. 40, 123–130 (2010).

89. Wiktor, J. & Wojciechowska, K. Differences in taxonomic composition of summer phytoplankton in two fjords of West Spitsbergen, Svalbard. Pol. Polar Res. 26, 259–268 (2005).

90. Włodarska-Kowalczuk, M., Szymelfenig, M., Kotwicki, L. & Warszawy, P. Macro- and meiobenthic fauna of the Yoldiabukta glacial bay (Isfjorden, Spitsbergen). Pol. Polar Res. 20, 367–386 (1999).

91. Włodarska-Kowalczuk, M. & Pearson, T. H. Soft-bottom macrobenthic faunal associations and factors affecting species distributions in an Arctic glacial fjord (Kongsfjord, Spitsbergen). Polar Biol. 27, 155–167 (2004).

92. Włodarska-Kowalczuk, M., Pearson, T. H. & Kendall, M. A. Benthic response to chronic natural physical disturbance by glacial sedimentation in an Arctic fjord. Mar. Ecol. Prog. Ser. 303, 31–41 (2005).

93. Włodarska-Kowalczuk, M. & Kedra, M. Surrogacy in natural patterns of benthic distribution and diversity: selected taxa versus lower taxonomic resolution. Mar. Ecol. Prog. Ser. 351, 53–63 (2007).

94. Włodarska-Kowalczuk, M. & Weslawski, J. M. Mesoscale spatial structures of soft-bottom macrozoobenthos communities: effects of physical control and impoverishment. Mar. Ecol. Prog. Ser. 356, 215–224 (2008).

95. Włodarska-Kowalczuk, M., Renaud, P. E., Weslawski, J. M., Cochrane, S. K. & Denisenko, S. G. Species diversity, functional complexity and rarity in Arctic fjordic versus open shelf benthic systems. Mar. Ecol. Prog. Ser. 463, 73–87 (2012).

96. Ziegler, A., Smith, C., Edwards, K. & Vernet, M. Glacial dropstones: islands enhancing seafloor species richness of benthic megafauna in West Antarctic Peninsula fjords. Mar. Ecol. Prog. Ser. 583, 1–14 (2017).

97. Zemko, K., Pabis, K., Siciński, J. & Błażewicz, M. Low abundance and high species richness: the structure of the soft-bottom isopod fauna of a West Antarctic glacial fjord. Polar Biol. 40, 2187–2199 (2017).

98. Battin, T. J., Wille, A., Sattler, B. & Psenner, R. Phylogenetic and functional heterogeneity of sediment biofilms along environmental gradients in a glacial stream. Appl. Environ. Microbiol. 67, 799–807 (2001).

99. Battin, T., Wille, A., Psenner, R. & Richter, A. Large-scale environmental controls on microbial biofilms in high-alpine streams. Biogeosciences 1, 159–171 (2004).

100. Blaen, P. J., Brown, L. E., Hannah, D. M. & Milner, A. M. Environmental drivers of macroinvertebrate communities in high Arctic rivers (Svalbard). Freshw. Biol. 59, 378–391 (2014).

101. Brittain, J. E. et al. The macroinvertebrate communities of two contrasting Norwegian glacial rivers in relation to environmental variables. Freshw. Biol. 46, 1723–1736 (2001).

102. Brown, L. E., Milner, A. M. & Hannah, D. M. Stability and persistence of alpine stream macroinvertebrate communities and the role of physicochemical habitat variables. Hydrobiologia 560, 159–173 (2006).

103. Brown, L. E., Hannah, D. M. & Milner, A. M. Vulnerability of alpine stream biodiversity to shrinking glaciers and snowpacks. Glob. Change Biol. 13, 958–966 (2007).

104. Brown, L. E., Milner, A. M. & Hannah, D. M. Predicting river ecosystem response to glacial meltwater dynamics: a case study of quantitative water sourcing and glaciality index approaches. Aquat. Sci. 72, 325–334 (2010).

105. Brown, L. E. & Milner, A. M. Rapid loss of glacial ice reveals stream community assembly processes. Glob. Change Biol. 18, 2195–2204 (2012).

106. Brown, L. E., Dickson, N., Carrivick, J. & Füreder, L. Alpine river ecosystem response to glacial and anthropogenic flow pulses. Freshw. Sci. 34, 1201–1215 (2015).

107. Brown, L. E. et al. Functional diversity and community assembly of river invertebrates show globally consistent responses to decreasing glacier cover. Nat. Ecol. Evol. 2, 325–333 (2018).

108. Burgherr, P. & Ward, J. Longitudinal and seasonal distribution patterns of the benthic fauna of an alpine glacial stream (Val Roseg, Swiss Alps). Freshw. Biol. 46, 1705–1721 (2001).

109. Cadbury, S. L., Milner, A. M. & Hannah, D. M. Hydroecology of a New Zealand glacier‐fed river: linking longitudinal zonation of physical habitat and macroinvertebrate communities. Ecohydrology 4, 520–531 (2011).

110. Castella, E. et al. Macrobenthic invertebrate richness and composition along a latitudinal gradient of European glacier‐fed streams. Freshw. Biol. 46, 1811–1831 (2001).

111. Cauvy-Fraunié, S., Espinosa, R., Andino, P., Dangles, O. & Jacobsen, D. Relationships between stream macroinvertebrate communities and new flood‐based indices of glacial influence. Freshw. Biol. 59, 1916–1925 (2014).

112. Di Lorenzo, T., Stoch, F. & Galassi, D. M. Incorporating the hyporheic zone within the river discontinuum: longitudinal patterns of subsurface copepod assemblages in an Alpine stream. Limnologica 43, 288–296 (2013).

113. Eisendle, U. Spatiotemporal distribution of free-living nematodes in glacial-fed stream reaches (Hohe Tauern, Eastern Alps, Austria). Arct. Antarct. Alp. Res. 40, 470–480 (2008).

114. Eisendle-Flöckner, U., Jersabek, C. D., Kirchmair, M., Hashold, K. & Traunspurger, W. Community patterns of the small riverine benthos within and between two contrasting glacier catchments. Ecol. Evol. 3, 2832–2844 (2013).

115. Elgmork, K. & Sæther, O. R. Distribution of Invertebrates in a High Mountain Brook in the Colorado Rocky Mountains (Colorado Univ. Press, 1970).

116. Esposito, R. et al. Antarctic climate cooling and response of diatoms in glacial meltwater streams. Geophys. Res. Lett. 33, L07406 (2006).

117. Finn, D. S., Rasanen, K. & Robinson, C. T. Physical and biological changes to a lengthening stream gradient following a decade of rapid glacial recession. Glob. Change Biol. 16, 3314–3326 (2010).

118. Finn, D. S., Khamis, K. & Milner, A. M. Loss of small glaciers will diminish beta diversity in Pyrenean streams at two levels of biological organization. Glob. Ecol. Biogeogr. 22, 40–51 (2013).

119. Fleming, S. W. Comparative analysis of glacial and nival streamflow regimes with implications for lotic habitat quantity and fish species richness. Riv. Res. Appl. 21, 363–379 (2005).

120. Freimann, R., Bürgmann, H., Findlay, S. E. & Robinson, C. T. Bacterial structures and ecosystem functions in glaciated floodplains: contemporary states and potential future shifts. ISME J. 7, 2361–2373 (2013).

121. Freimann, R., Bürgmann, H., Findlay, S. E. & Robinson, C. T. Spatio-temporal patterns of major bacterial groups in Alpine waters. PLoS ONE 9, e113524 (2014).

122. Friberg, N., Milner, A. M., Svendsen, L. M., Lindegaard, C. & Larsen, S. E. Macroinvertebrate stream communities along regional and physico‐chemical gradients in Western Greenland. Freshw. Biol. 46, 1753–1764 (2001).

123. Füreder, L., Schütz, C., Wallinger, M. & Burger, R. Physico‐chemistry and aquatic insects of a glacier‐fed and a spring‐fed alpine stream. Freshw. Biol. 46, 1673–1690 (2001).

124. Füreder, L. et al. Reference conditions of alpine streams: physical habitat and ecology. Water Air Soil Poll. 2, 275–294 (2002).

125. Füreder, L. Life at the edge: habitat condition and bottom fauna of Alpine running waters. Int. Rev. Hydrobiol. 92, 491–513 (2007).

126. Gesierich, D. & Rott, E. Benthic algae and mosses from aquatic habitats in the catchment of a glacial stream (Rotmoos, Ötztal, Austria). Ber. Naturwiss.-med. Ver. Innsb. 91, 7–42 (2004).

127. Gesierich, D. & Rott, E. Is diatom richness responding to catchment glaciation? A case study from Canadian headwater streams. J. Limnol. 71, 72–83 (2012).

128. Gislason, G. M., Adalsteinsson, H., Olafsson, J. S. & Hansen, I. Invertebrate communities of glacial and alpine rivers in the central highlands of Iceland. Verh. Int. Ver. Theor. Angew. Limnol. 27, 1602–1606 (2001).

129. Gíslason, G. M., Hansen, I., Ólafsson, J. S. & Svavarsdóttir, K. Longitudinal changes in macroinvertebrate assemblages along a glacial river system in central Iceland. Freshw. Biol. 46, 1737–1751 (2001).

130. Hamerlik, L. & Jacobsen, D. Chironomid (Diptera) distribution and diversity in Tibetan streams with different glacial influence. Insect Conserv. Divers. 5, 319–326 (2012).

131. Hansen, I., Gíslason, G. M. & Olafsson, J. S. Diatoms in glacial and alpine rivers in Central Iceland. Verh. Int. Ver. Theor. Angew. Limnol. 29, 1271–1274 (2006).

132. Hieber, M., Robinson, C. T., Rushforth, S. R. & Uehlinger, U. Algal communities associated with different alpine stream types. Arct. Antarct. Alp. Res. 33, 447–456 (2001).

133. Hieber, M., Robinson, C. T. & Uehlinger, U. Seasonal and diel patterns of invertebrate drift in different alpine stream types. Freshw. Biol. 48, 1078–1092 (2003).

134. Hieber, M., Robinson, C. T., Uehlinger, U. & Ward, J. A comparison of benthic macroinvertebrate assemblages among different types of alpine streams. Freshw. Biol. 50, 2087–2100 (2005).

135. Howard‐Williams, C., Vincent, C. L., Broady, P. A. & Vincent, W. F. Antarctic stream ecosystems: variability in environmental properties and algal community structure. Int. Rev. Ges. Hydrobiol. Hydrogr. 71, 511–544 (1986).

136. Huryn, A. D. et al. Landscape heterogeneity and the biodiversity of Arctic stream communities: a habitat template analysis. Can. J. Fish. Aquat. Sci. 62, 1905–1919 (2005).

137. Hylander, S. et al. Climate-induced input of turbid glacial meltwater affects vertical distribution and community composition of phyto- and zooplankton. J. Plankton Res. 33, 1239–1248 (2011).

138. Ilg, C. & Castella, E. Patterns of macroinvertebrate traits along three glacial stream continuums. Freshw. Biol. 51, 840–853 (2006).

139. Jacobsen, D. et al. Longitudinal zonation of macroinvertebrates in an Ecuadorian glacier-fed stream: do tropical glacial systems fit the temperate model? Freshw. Biol. 55, 1234–1248 (2010).

140. Khamis, K., Hannah, D., Brown, L., Tiberti, R. & Milner, A. The use of invertebrates as indicators of environmental change in alpine rivers and lakes. Sci. Total Environ. 493, 1242–1254 (2014).

141. Khamis, K., Brown, L. E., Hannah, D. M. & Milner, A. M. Glacier–groundwater stress gradients control alpine river biodiversity. Ecohydrology 9, 1263–1275 (2016).

142. Knispel, S. & Castella, E. Disruption of a longitudinal pattern in environmental factors and benthic fauna by a glacial tributary. Freshw. Biol. 48, 604–618 (2003).

143. Kuhn, J. et al. Spatial variability in macroinvertebrate assemblages along and among neighbouring equatorial glacier‐fed streams. Freshw. Biol. 56, 2226–2244 (2011).

144. Lencioni, V., Maiolini, B. & Rossaro, B. The kryal and rhithral Chironomid community in the Carè Alto system (Italian central-eastern Alps). Verh. Int. Ver. Theor. Angew. Limnol. 27, 711–715 (2000).

145. Lencioni, V. & Rossaro, B. Microdistribution of chironomids (Diptera: Chironomidae) in Alpine streams: an autoecological perspective. Hydrobiologia 533, 61–76 (2005).

146. Lencioni, V. Glacial influence and stream macroinvertebrate biodiversity under climate change: lessons from the Southern Alps. Sci. Total Environ. 622, 563–575 (2018).

147. Lods-Crozet, B. et al. Macroinvertebrate community structure in relation to environmental variables in a Swiss glacial stream. Freshw. Biol. 46, 1641–1661 (2001).

148. Lods-Crozet, B. et al. Chironomid (Diptera: Chironomidae) communities in six European glacier‐fed streams. Freshw. Biol. 46, 1791–1809 (2001).

149. Lods-Crozet, B., Lencioni, V., Brittain, J. E., Marziali, L. & Rossaro, B. Contrasting chironomid assemblages in two high Arctic streams on Svalbard. Fund. Appl. Limnol. Arch. Hydrobiol. 170, 211–222 (2007).

150. Maiolini, B. & Lencioni, V. Longitudinal distribution of macroinvertebrate assemblages in a glacially influenced stream system in the Italian Alps. Freshw. Biol. 46, 1625–1639 (2001).

151. Malard, F., Galassi, D., Lafont, M., Dolédec, S. & Ward, J. Longitudinal patterns of invertebrates in the hyporheic zone of a glacial river. Freshw. Biol. 48, 1709–1725 (2003).

152. Milner, A. M. Colonization and ecological development of new streams in Glacier Bay National Park, Alaska. Freshw. Biol. 18, 53–70 (1987).

153. Milner, A. M. et al. Colonization and development of stream communities across a 200-year gradient in Glacier Bay National Park, Alaska, USA. Can. J. Fish. Aquat. Sci. 57, 2319–2335 (2000).

154. Milner, A. M. et al. Evolution of a stream ecosystem in recently deglaciated terrain. Ecology 92, 1924–1935 (2011).

155. Miserendino, M. L. et al. Biotic diversity of benthic macroinvertebrates at contrasting glacier-fed systems in Patagonia Mountains: the role of environmental heterogeneity facing global warming. Sci. Total Environ. 622, 152–163 (2018).

156. Molina, J. M. Diversidad de Grupos Functionales Troficos de Macroinvertebrados, en los Rios de la Cordillera Real y las Serranias Altiplanicas de la Hidroecoregion Altoandina. MS thesis, Universidad Mayor de San Andrés (2013).

157. Murakami, T. et al. Limnological features of glacier-fed rivers in the Southern Tibetan Plateau, China. Limnology 13, 301–307 (2012).

158. Olafsson, J. S., Mar, G. G. & Adalsteinsson, H. Chironomids in glacial and non-glacial rivers in Iceland: a comparative study. Verh. Int. Ver. Theor. Angew. Limnol. 27, 720–726 (2001).

159. Parker, S. M. & Huryn, A. D. Effects of natural disturbance on stream communities: a habitat template analysis of arctic headwater streams. Freshw. Biol. 56, 1342–1357 (2011).

160. Peter, H. & Sommaruga, R. Shifts in diversity and function of lake bacterial communities upon glacier retreat. ISME J. 10, 1545–1554 (2016).

161. Quenta, E. et al. Direct and indirect effects of glaciers on aquatic biodiversity in high Andean peatlands. Glob. Change Biol. 22, 3196–3205 (2016).

162. Ren, Z., Gao, H. & Elser, J. J. Longitudinal variation of microbial communities in benthic biofilms and association with hydrological and physicochemical conditions in glacier-fed streams. Freshw. Sci. 36, 479–490 (2017).

163. Robertson, A. & Milner, A. Meiobenthic arthropod communities in new streams in Glacier Bay National Park, Alaska. Hydrobiologia 397, 197–209 (1999).

164. Robertson, A. & Milner, A. The influence of stream age and environmental variables in structuring meiofaunal assemblages in recently deglaciated streams. Limnol. Oceanogr. 51, 1454–1465 (2006).

165. Robinson, C. T., Gessner, M. O., Callies, K. A., Jolidon, C. & Ward, J. V. Larch needle breakdown in contrasting streams of an alpine glacial floodplain. J. N. Am. Benthol. Soc. 19, 250–262 (2000).

166. Robinson, C., Uehlinger, U. & Hieber, M. Spatio‐temporal variation in macroinvertebrate assemblages of glacial streams in the Swiss Alps. Freshw. Biol. 46, 1663–1672 (2001).

167. Robinson, C. & Jolidon, C. Leaf breakdown and the ecosystem functioning of alpine streams. J. N. Am. Benthol. Soc. 24, 495–507 (2005).

168. Robinson, C. T. & Kawecka, B. Benthic diatoms of an Alpine stream/lake network in Switzerland. Aquat. Sci. 67, 492–506 (2005).

169. Robinson, C. T., Hieber, M., Wenzelides, V. & Lods-Crozet, B. Macroinvertebrate assemblages of a high elevation stream/lake network with an emphasis on the Chironomidae. Fund. Appl. Limnol. Arch. Hydrobiol. 169, 25–36 (2007).

170. Rott, E., Cantonati, M., Füreder, L. & Pfister, P. Benthic algae in high altitude streams of the Alps—a neglected component of the aquatic biota. Hydrobiologia 562, 195–216 (2006).

171. Sheath, R. G. & Müller, K. M. Distribution of stream macroalgae in four high Arctic drainage basins. Arctic 50, 355–364 (1997).

172. Slemmons, K. E. & Saros, J. E. Implications of nitrogen‐rich glacial meltwater for phytoplankton diversity and productivity in alpine lakes. Limnol. Oceanogr. 57, 1651–1663 (2012).

173. Snook, D. L. & Milner, A. M. Biological traits of macroinvertebrates and hydraulic conditions in a glacier-fed catchment (French Pyrenees). Arch. Hydrobiol. 153, 245–271 (2002).

174. Thompson, C., David, E., Freestone, M. & Robinson, C. Ecological patterns along two alpine glacial streams in the Fitzpatrick Wilderness, Wind River Range, USA. West. N. Am. Nat. 73, 137–147 (2013).

175. Turner, K. L., Matthews, R. A. & Rawhouser, A. K. Benthic macroinvertebrate assemblages in kryal and rhithral lake outlet streams in the North Cascade Mountains. Northwest Sci. 90, 206–227 (2016).

176. Wesener, M. D. et al. Hyporheic and benthic macroinvertebrate communities in glacial, clearwater, and brownwater streams in Alaska. Pan-Pac. Entomol. 87, 145–160 (2011).

177. Albrecht, M., Riesen, M. & Schmid, B. Plant–pollinator network assembly along the chronosequence of a glacier foreland. Oikos 119, 1610–1624 (2010).

178. Alfredsen, G. & Høiland, K. Succession of terrestrial macrofungi along a deglaciation gradient at Glacier Blåisen, South Norway. Nord. J. Bot. 21, 19–37 (2001).

179. Andreis, C., Caccianiga, M. & Cerabolini, B. Vegetation and environmental factors during primary succession on glacier forelands: some outlines from the Italian Alps. Plant Biosyst. 135, 295–310 (2001).

180. Bajerski, F. & Wagner, D. Bacterial succession in Antarctic soils of two glacier forefields on Larsemann Hills, East Antarctica. FEMS Microbiol. Ecol. 85, 128–142 (2013).

181. Bárcena, T. G., Yde, J. C. & Finster, K. W. Methane flux and high-affinity methanotrophic diversity along the chronosequence of a receding glacier in Greenland. Ann. Glaciol. 51, 23–31 (2010).

182. Birks, H. J. The present flora and vegetation of the moraines of the Klutlan Glacier, Yukon Territory, Canada: a study in plant succession. Quat. Res. 14, 60–86 (1980).

183. Blaalid, R. et al. Changes in the root-associated fungal communities along a primary succession gradient analysed by 454 pyrosequencing. Mol. Ecol. 21, 1897–1908 (2012).

184. Bradley, J. A. et al. Microbial dynamics in a High Arctic glacier forefield: a combined field, laboratory, and modelling approach. Biogeosciences 13, 5677–5696 (2016).

185. Bråten, A. T. et al. Primary succession of surface active beetles and spiders in an alpine glacier foreland, central south Norway. Arct. Antarct. Alp. Res. 44, 2–15 (2012).

186. Burga, C. A. Vegetation development on the glacier forefield Morteratsch (Switzerland). Appl. Veg. Sci. 2, 17–24 (1999).

187. Carlson, M. L., Flagstad, L. A., Gillet, F. & Mitchell, E. A. Community development along a proglacial chronosequence: are above‐ground and below‐ground community structure controlled more by biotic than abiotic factors? J. Ecol. 98, 1084–1095 (2010).

188. Conen, F., Yakutin, M., Zumbrunn, T. & Leifeld, J. Organic carbon and microbial biomass in two soil development chronosequences following glacial retreat. Eur. J. Soil Sci. 58, 758–762 (2007).

189. Deiglmayr, K., Philippot, L., Tscherko, D. & Kandeler, E. Microbial succession of nitrate‐reducing bacteria in the rhizosphere of Poa alpina across a glacier foreland in the Central Alps. Environ. Microbiol. 8, 1600–1612 (2006).

190. Dong, K. et al. Soil fungal community development in a high Arctic glacier foreland follows a directional replacement model, with a mid-successional diversity maximum. Sci. Rep. 6, 26360 (2016).

191. Duc, L., Noll, M., Meier, B. E., Bürgmann, H. & Zeyer, J. High diversity of diazotrophs in the forefield of a receding alpine glacier. Microb. Ecol. 57, 179–190 (2009).

192. Erschbamer, B. & Mayer, R. Can successional species groups be discriminated based on their life history traits? A study from a glacier foreland in the Central Alps. Plant Ecol. Divers. 4, 341–351 (2011).

193. Esperschütz, J. et al. Microbial food web dynamics along a soil chronosequence of a glacier forefield. Biogeosciences 8, 3283–3294 (2011).

194. Fernández-Martínez, M. A. et al. Microbial succession dynamics along glacier forefield chronosequences in Tierra del Fuego (Chile). Polar Biol. 40, 1939–1957 (2017).

195. Franzen, M. & Dieker, P. The influence of terrain age and altitude on the arthropod communities found on recently deglaciated terrain. Curr. Zool. 60, 203–220 (2014).

196. Frenot, Y., Gloaguen, J., Cannavacciuolo, M. & Bellido, A. Primary succession on glacier forelands in the subantarctic Kerguelen Islands. J. Veg. Sci. 9, 75–84 (1998).

197. Frey, B., Bühler, L., Schmutz, S., Zumsteg, A. & Furrer, G. Molecular characterization of phototrophic microorganisms in the forefield of a receding glacier in the Swiss Alps. Environ. Res. Lett. 8, 015033 (2013).

198. Fujiyoshi, M. et al. Successional changes in ectomycorrhizal fungi associated with the polar willow Salix polaris in a deglaciated area in the High Arctic, Svalbard. Polar Biol. 34, 667–673 (2011).

199. Garibotti, I. A., Pissolito, C. I. & Villalba, R. Vegetation development on deglaciated rock outcrops from Glaciar Frias, Argentina. Arct. Antarct. Alp. Res. 43, 35–45 (2011).

200. Gereben-Krenn, B.-A., Krenn, H. W. & Strodl, M. A. Initial colonization of new terrain in an Alpine glacier foreland by carabid beetles (Carabidae, Coleoptera). Arct. Antarct. Alp. Res. 43, 397–403 (2011).

201. Gobbi, M., Fontaneto, D. & De Bernardi, F. Influence of climate changes on animal communities in space and time: the case of spider assemblages along an alpine glacier foreland. Glob. Change Biol. 12, 1985–1992 (2006).

202. Gobbi, M., Bernardi, F. D., Pelfini, M., Rossaro, B. & Brandmayr, P. Epigean arthropod succession along a 154-year glacier foreland chronosequence in the Forni Valley (Central Italian Alps). Arct. Antarct. Alp. Res. 38, 357–362 (2006).

203. Gobbi, M. et al. Plant adaptive responses during primary succession are associated with functional adaptations in ground beetles on deglaciated terrain. Commun. Ecol. 11, 223–231 (2010).

204. Gobbi, M. et al. Life in harsh environments: carabid and spider trait types and functional diversity on a debris‐covered glacier and along its foreland. Ecol. Entomol. 42, 838–848 (2017).

205. Górniak, D., Marszałek, H., Kwaśniak-Kominek, M., Rzepa, G. & Manecki, M. Soil formation and initial microbiological activity on a foreland of an Arctic glacier (SW Svalbard). Appl. Soil Ecol. 114, 34–44 (2017).

206. Gryziak, G. Colonization by mites of glacier-free areas in King George Island, Antarctica. Pesq. Agropec. Bras. 44, 891–895 (2009).

207. Hågvar, S., Solhøy, T. & Mong, C. E. Primary succession of soil mites (Acari) in a Norwegian glacier foreland, with emphasis on oribatid species. Arct. Antarct. Alp. Res. 41, 219–227 (2009).

208. Hågvar, S. Primary succession of springtails (Collembola) in a Norwegian glacier foreland. Arct. Antarct. Alp. Res. 42, 422–429 (2010).

209. Hodkinson, I. D., Coulson, S. J. & Webb, N. R. Community assembly along proglacial chronosequences in the high Arctic: vegetation and soil development in north‐west Svalbard. J. Ecol. 91, 651–663 (2003).

210. Hodkinson, I. D., Coulson, S. J. & Webb, N. R. Invertebrate community assembly along proglacial chronosequences in the high Arctic. J. Anim. Ecol. 73, 556–568 (2004).

211. Ilieva-Makulec, K. & Gryziak, G. Response of soil nematodes to climate-induced melting of Antarctic glaciers. Pol. J. Ecol. 57, 811–816 (2009).

212. Ingimarsdóttir, M. et al. Primary assembly of soil communities: disentangling the effect of dispersal and local environment. Oecologia 170, 745–754 (2012).

213. Insam, H. et al. Soil microbiota along Ayoloco glacier retreat area of Iztaccíhuatl volcano. Mex. Catena 153, 83–88 (2017).

214. Jones, G. A. & Henry, G. H. Primary plant succession on recently deglaciated terrain in the Canadian High Arctic. J. Biogeogr. 30, 277–296 (2003).

215. Jumpponen, A., Trappe, J. M. & Cázares, E. Occurrence of ectomycorrhizal fungi on the forefront of retreating Lyman Glacier (Washington, USA) in relation to time since deglaciation. Mycorrhiza 12, 43–49 (2002).

216. Jumpponen, A., Brown, S. P., Trappe, J. M., Cázares, E. & Strömmer, R. Twenty years of research on fungal–plant interactions on Lyman Glacier forefront—lessons learned and questions yet unanswered. Fungal Ecol. 5, 430–442 (2012).

217. Kaštovská, K., Elster, J., Stibal, M. & Šantrůčková, H. Microbial assemblages in soil microbial succession after glacial retreat in Svalbard (High Arctic). Microb. Ecol. 50, 396–407 (2005).

218. Kaufmann, R. Invertebrate succession on an alpine glacier foreland. Ecology 82, 2261–2278 (2001).

219. Kim, M., Jung, J. Y., Laffly, D., Kwon, H. Y. & Lee, Y. K. Shifts in bacterial community structure during succession in a glacier foreland of the High Arctic. FEMS Microbiol. Ecol. 93, fiw213 (2017).

220. Lazzaro, A., Risse-Buhl, U. & Brankatschk, R. Molecular and morphological snapshot characterisation of the protist communities in contrasting Alpine glacier forefields. Acta Protozool. 54, 143–154 (2015).

221. Liu, J. et al. Diversity and succession of autotrophic microbial community in high-elevation soils along deglaciation chronosequence. FEMS Microbiol. Ecol. 92, fiw160 (2016).

222. Mateos-Rivera, A. et al. The effect of temperature change on the microbial diversity and community structure along the chronosequence of the sub-arctic glacier forefield of Styggedalsbreen (Norway). FEMS Microbiol. Ecol. 92, fnw038 (2016).

223. Mizuno, K. Succession processes of alpine vegetation in response to glacial fluctuations of Tyndall Glacier, Mt. Kenya, Kenya. Arct. Alp. Res. 30, 340–348 (1998).

224. Mizuno, K. & Fujita, T. Vegetation succession on Mt. Kenya in relation to glacial fluctuation and global warming. J. Veg. Sci. 25, 559–570 (2014).

225. Müller, E., Eidesen, P. B., Ehrich, D. & Alsos, I. G. Frequency of local, regional, and long-distance dispersal of diploid and tetraploid Saxifraga oppositifolia (Saxifragaceae) to Arctic glacier forelands. Am. J. Bot. 99, 459–471 (2012).

226. Nascimbene, J., Mayrhofer, H., Dainese, M. & Bilovitz, P. O. Assembly patterns of soil‐dwelling lichens after glacier retreat in the European Alps. J. Biogeogr. 44, 1393–1404 (2017).

227. Nemergut, D. R. et al. Microbial community succession in an unvegetated, recently deglaciated soil. Microbiol. Ecol. 53, 110–122 (2007).

228. Pérez, C. A. et al. Ecosystem development in short‐term postglacial chronosequences: N and P limitation in glacier forelands from Santa Inés Island, Magellan Strait. Austral Ecol. 39, 288–303 (2014).

229. Philippot, L., Tscherko, D., Bru, D. & Kandeler, E. Distribution of high bacterial taxa across the chronosequence of two alpine glacier forelands. Microbiol. Ecol. 61, 303–312 (2011).

230. Raffl, C., Mallaun, M., Mayer, R. & Erschbamer, B. Vegetation succession pattern and diversity changes in a glacier valley, Central Alps, Austria. Arct. Antarct. Alp. Res. 38, 421–428 (2006).

231. Reiners, W. A., Worley, I. A. & Lawrence, D. B. Plant diversity in a chronosequence at Glacier Bay, Alaska. Ecology 52, 55–69 (1971).

232. Rime, T. et al. Vertical distribution of the soil microbiota along a successional gradient in a glacier forefield. Mol. Ecol. 24, 1091–1108 (2015).

233. Robbins, J. A. & Matthews, J. A. Pioneer vegetation on glacier forelands in southern Norway: emerging communities? J. Veg. Sci. 20, 889–902 (2009).

234. Schlag, R. N. & Erschbamer, B. Germination and establishment of seedlings on a glacier foreland in the Central Alps, Austria. Arct. Antarct. Alp. Res. 32, 270–277 (2000).

235. Schütte, U. M. et al. Bacterial succession in a glacier foreland of the High Arctic. ISME J. 3, 1258–1268 (2009).

236. Sigler, W., Crivii, S. & Zeyer, J. Bacterial succession in glacial forefield soils characterized by community structure, activity and opportunistic growth dynamics. Microbiol. Ecol. 44, 306–316 (2002).

237. Sigler, W. & Zeyer, J. Microbial diversity and activity along the forefields of two receding glaciers. Microbiol. Ecol. 43, 397–407 (2002).

238. Simmons, B. et al. Long-term experimental warming reduces soil nematode populations in the McMurdo Dry Valleys, Antarctica. Soil Biol. Biochem. 41, 2052–2060 (2009).

239. Srinivas, T. et al. Comparison of bacterial diversity in proglacial soil from Kafni Glacier, Himalayan Mountain ranges, India, with the bacterial diversity of other glaciers in the world. Extremophiles 15, 673–690 (2011).

240. Stöcklin, J. & Bäumler, E. Seed rain, seedling establishment and clonal growth strategies on a glacier foreland. J. Veg. Sci. 7, 45–56 (1996).

241. Tian, J. et al. Ecological succession pattern of fungal community in soil along a retreating glacier. Front. Microbiol. 8, 1028 (2017).

242. Trowbridge, J. & Jumpponen, A. Fungal colonization of shrub willow roots at the forefront of a receding glacier. Mycorrhiza 14, 283–293 (2004).

243. Tscherko, D., Rustemeier, J., Richter, A., Wanek, W. & Kandeler, E. Functional diversity of the soil microflora in primary succession across two glacier forelands in the Central Alps. Eur. J. Soil Sci. 54, 685–696 (2003).

244. Tscherko, D., Hammesfahr, U., Zeltner, G., Kandeler, E. & Böcker, R. Plant succession and rhizosphere microbial communities in a recently deglaciated alpine terrain. Basic Appl. Ecol. 6, 367–383 (2005).

245. Vater, A. E. Insect and arachnid colonization on the Storbreen glacier foreland, Jotunheimen, Norway: persistence of taxa suggests an alternative model of succession. Holocene 22, 1123–1133 (2012).

246. Vater, A. E. & Matthews, J. A. Testing the ‘addition and persistence model’ of invertebrate succession in a subalpine glacier-foreland chronosequence: Fåbergstølsbreen, southern Norway. Holocene 23, 1151–1162 (2013).

247. Vater, A. E. & Matthews, J. A. Succession of pitfall-trapped insects and arachnids on eight Norwegian glacier forelands along an altitudinal gradient: patterns and models. Holocene 25, 108–129 (2015).

248. Wu, X. et al. Bacterial diversity in the foreland of the Tianshan No. 1 glacier, China. Environ. Res. Lett. 7, 014038 (2012).

249. Zimmer, A. et al. Time lag between glacial retreat and upward migration alters tropical alpine communities. Perspect. Plant Ecol. 30, 89–102 (2017).

250. Zumsteg, A. et al. Bacterial, archaeal and fungal succession in the forefield of a receding glacier. Microbiol. Ecol. 63, 552–564 (2012).

251. Lajeunesse, M. J., Koricheva, J., Gurevitch, J. & Mengersen, K. in Handbook of Meta-analysis in Ecology and Evolution 195–206 (2013).

252. Hedges, L. V. & Olkin, I. Statistical Methods for Meta-analysis (Academic Press, 1985).

253. Borenstein, M., Hedges, L. V., Higgins, J. P. & Rothstein, H. R. Introduction to Meta-analysis (John Wiley & Sons, 2009).

254. Gardner, A. S. et al. A reconciled estimate of glacier contributions to sea level rise: 2003 to 2009. Science 340, 852–857 (2013).

255. Nakagawa, S., Noble, D. W., Senior, A. M. & Lagisz, M. Meta-evaluation of meta-analysis: ten appraisal questions for biologists. BMC Biol. 15, 18 (2017).

256. Nakagawa, S. & Santos, E. S. Methodological issues and advances in biological meta-analysis. Evol. Ecol. 26, 1253–1274 (2012).

257. Konstantopoulos, S. Fixed effects and variance components estimation in three‐level meta‐analysis. Res. Synth. Methods 2, 61–76 (2011).

258. Finlay, B. J. Global dispersal of free-living microbial eukaryote species. Science 296, 1061–1063 (2002).

259. Viechtbauer, W. Conducting meta-analyses in R with the metafor package. J. Stat. Softw. 36, 1–48 (2010).

260. Peters, J. L., Sutton, A. J., Jones, D. R., Abrams, K. R. & Rushton, L. Contour-enhanced meta-analysis funnel plots help distinguish publication bias from other causes of asymmetry. J. Clin. Epidemiol. 61, 991–996 (2008).

261. Egger, M., Smith, G. D., Schneider, M. & Minder, C. Bias in meta-analysis detected by a simple, graphical test. Br. Med. J. 315, 629–634 (1997).

262. Pfeffer, W. T. et al. The Randolph Glacier Inventory: a globally complete inventory of glaciers. J. Glaciol. 60, 537–552 (2014).