Seventy-five mosses and liverworts found frozen with the late Neolithic Tyrolean Iceman: Origins, taphonomy and the Iceman’s last journey

Autoři: James H. Dickson aff001;  Klaus D. Oeggl aff002;  Werner Kofler aff002;  Wolfgang K. Hofbauer aff003;  Ronald Porley aff004;  Gordon P. Rothero aff001;  Alexandra Schmidl aff002;  Andreas G. Heiss aff002
Působiště autorů: Institute of Biodiversity, Animal Health & Comparative Medicine, University of Glasgow, Glasgow, Scotland, United Kingdom aff001;  Institut für Botanik, University of Innsbruck, Innsbruck, Austria aff002;  Franhofer Institut Bauphysik, Valley, Germany aff003;  English Nature, Crookham Common, United Kingdom aff004
Vyšlo v časopise: PLoS ONE 14(10)
Kategorie: Research Article
doi: 10.1371/journal.pone.0223752


The Iceman site is unique in the bryology of the Quaternary. Only 21 bryophytes (mosses and liverworts) grow now in the immediate vicinity of the 5,300 year old Iceman discovery site at 3,210m above sea level in the Ötztal Alps, Italy. By contrast 75 or more species including at least ten liverworts were recovered as subfossils frozen in, on and around the Iceman from before, at and after his time. About two thirds of the species grow in the nival zone (above 3,000m above sea level) now while about one third do not. A large part of this third can be explained by the Iceman having both deliberately and inadvertently carried bryophytes during his last, fatal journey. Multivariate analyses (PCA, RDA) provide a variety of explanations for the arrivals of the bryophytes in the rocky hollow where the mummy was discovered. This is well into the nival zone of perennial snow and ice with a very sparse, non-woody flora and very low vegetation cover. Apart from the crucial anthropochory (extra-local plants), both hydrochory (local species) and zoochory (by wild game such as ibex of both local and extra-local species) have been important. Anemochory of mainly local species was of lesser importance and of extra-local species probably of little or no importance. The mosses Neckera complanata and several other ecologically similar species as well as a species of Sphagnum (bogmoss) strongly support the claim that the Iceman, took northwards up Schnalstal, South Tyrol, as the route of the last journey. A different species of bogmoss, taken from his colon is another indication the Iceman’s presence at low altitude south of Schnalstal during his last hours when he was first high up, low down and finally at over 3,000m.

Klíčová slova:

Equipment – Grasses – Principal component analysis – Sediment – Bryology – Nonvascular plants – Mosses – Subfossils


1. Kutschera W, Patzelt G, Wild EM, Haas-Jettmar B, Kofler W, Lippert A, et al. Evidence for Early Human Presence At High Altitudes in the Otztal Alps (Austria/Italy). Radiocarbon. 2014;56(3): 923–947.

2. Wierer U, Arrighi S, Bertola S, Kaufmann G, Baumgarten B, Pedrotti A, et al. The Iceman's lithic toolkit: Raw material, technology, typology and use. PLOS One. 2018;13(6): e0198292. doi: 10.1371/journal.pone.0198292 29924811

3. Pernter P, Gostner P, Vigl EE, Rühli FJ. Radiologic proof for the Iceman's cause of death (ca. 5’300 BP). J Archaeol Sci. 2007;34(11): 1784–1786.

4. Egg M, Spindler K. Kleidung und Ausrüstung der kupferzeitlichen Gletschermumie aus den Ötztaler Alpen. Mainz: Verl. des Römisch-German. Zentralmuseums; 2009.

5. Dickson JH. Ancient ice mummies. Stroud: The History Press; 2011.

6. Rollo F, Ubaldi M, Ermini L, Marota I. Ötzi's last meals: DNA analysis of the intestinal content of the Neolithic glacier mummy from the Alps. Proc Natl Acad Sci U S A. 2002;99(20): 12594–12599. doi: 10.1073/pnas.192184599 12244211

7. Maixner F, Turaev D, Cazenave-Gassiot A, Janko M, Krause-Kyora B, Hoopmann MR, et al. The Iceman’s Last Meal Consisted of Fat, Wild Meat, and Cereals. Curr Biol. 2018;28(14): 2348–55.e9. doi: 10.1016/j.cub.2018.05.067 30017480

8. Oeggl K, Kofler W, Schmidl A. War Ötzi wirklich ein Hirte?. Berichte der Reinhold-Tüxen-Gesellschaft. 2005; 17: 137–149.

9. Oeggl K. The significance of the Tyrolean Iceman for the archaeobotany of Central Europe. Veg Hist Archaeobot. 2009;18(1): 1–11.

10. Festi D, Putzer A, Oeggl K. Mid and late Holocene land-use changes in the Otztal Alps, territory of the Neolithic Iceman "Otzi". Quat Int. 2014;353: 17–33.

11. Reitmaier T. Form follows function—a new interpretation of the so-called stone disc with tassel of the South Tyrolean ice man. Mainz: Publisher of the Roman-Germanic Central Museum 2014. pp. 29–40.

12. Artioli G, Angelini I, Kaufmann G, Canovaro C, Dal Sasso G, Villa IM. Long-distance connections in the Copper Age: New evidence from the Alpine Iceman's copper axe. PLoS One. 2017;12(7): e0179263. doi: 10.1371/journal.pone.0179263 28678801

13. Bortenschlager S, Oeggl K. The Iceman and his natural environment. The Man in the Ice Vol. 4. New York: Springer; 2000.

14. Heiss AG, Oeggl K. The plant macro-remains from the Iceman site (Tisenjoch, Italian—Austrian border, eastern Alps): new results on the glacier mummy's environment. Veg Hist Archaeobot. 2009;18(1): 23–35.

15. Oeggl K, Kofler W, Schmidl A, Dickson JH, Egarter-Vigl E, Gaber O. The reconstruction of the last itinerary of “Ötzi”, the Neolithic Iceman, by pollen analyses from sequentially sampled gut extracts. Quat Sci Rev. 2007;26(7): 853–61.

16. Dickson JH. The moss from the Tyrolean Iceman's colon. J Bryol 1997;19: 449–51.

17. Dickson JH. Bryology and the Iceman: Chorology, Ecology and Ethnobotany of the Mosses Neckera complanata Hedw. and N. crispa Hedw. The man in the ice Volume 4: The iceman and his natural environment: palaeobotanical results. Wien: Springer; 2000. pp. 77–88.

18. Dickson JH. Low to moderate altitude bryophytes at the Iceman site and their significance. Bolzano: South Tyrol Museum of Archeology; 2003. pp. 27–34.

19. Dickson JH. Bogmoss in the Iceman's stomach. The Bryological Times 2013;137: 28–30.

20. Dickson JH, Bortenschlager S, Oeggl K, Porley R, McMullen A. Mosses and the Tyrolean Iceman's Southern Provenance. Proc R Soc Lond B Biol Sci. 1996;263(1370): 567–571.

21. Dickson JH, Hofbauer W, Kofler W, Oeggl K, Platzgummer J. How to find the bogmoss, Sphagnum imbricatum s.l., in South Tyrol, Italy: Microscopically examine the Iceman's colon contents. Veg Hist Archaeobot. 2005;14(3): 207–210.

22. Dickson JH, Hofbauer W, Porley R, Schmidl A, Kofler W, Oeggl K. Six mosses from the Tyrolean Iceman's alimentary tract and their significance for his ethnobotany and the events of his last days. Veg Hist Archaeobot. 2009;18(1): 13–22.

23. Gottfried M, Pauli H, Reiter K, Grabherr G. Potential Effects of Climatic Change on Alpine and Nival Plants in the Alps. In: Korner C, Spehn EM, editors. Mountain Biodiversity: A Global Assessment. Boca Raton (FL): CRC Press; 2002. pp. 213–224.

24. Heinken T. Dispersal of plants by a dog in a deciduous forest. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie. 2000;122: 449–468.

25. Heinken T, Lees R, Raudnitschka D, Runge S. Epizoochorous dispersal of bryophyte stem fragments by roe deer (Capreolus capreolus) and wild boar (Sus scrofa). J Bryol. 2001;23: 293–300.

26. Pauliuk F, Muller J, Heinken T. Bryophyte dispersal by sheep on dry grassland. Nova Hedwigia. 2011;92(3–4): 327–41.

27. Lewis LR, Behling E, Gousse H, Qian E, Elphick CS, Lamarre JF, et al. First evidence of bryophyte diaspores in the plumage of transequatorial migrant birds. PEERJ. 2014;2: e424. doi: 10.7717/peerj.424 24949241

28. Wilkinson DM, Lovas-Kiss A, Callaghan DA, Green AJ. Endozoochory of large bryophyte fragments by waterbirds. Cryptogamie Bryologie. 2017;38(2): 223–238.

29. Miller NG, Howe-Ambrose LJ. Growth in Culture of Wind-Blown Bryophyte Gametophyte Fragments from Arctic Canada. Bryologist. 1976;79(1): 55–63.

30. Miller NG. Fossil evidence of the dispersal and establishment of mosses as gametophyte fragments. Monographs in systematic botany from the Missouri Botanical Garden (USA). 1985;11: 71–85.

31. McDaniel SF, Miller NG. Winter Dispersal of Bryophyte Fragments in the Adirondack Mountains, New York. Bryologist. 2000;103(3): 592–600.

32. Lippert A. Die erste archäologische Nachuntersuchung am Tisenjoch. Der Mann im Eis: Band 1. 187. Innsbruck: Publications of the University of Innsbruck; 1992. pp. 245–253.

33. Lippert A, Spindler K. Die Auffindung einer frühbronzezeitlichen Gletschermumie am Hauslabjoch in den Ötztaler Alpen (Gem. Schnals). Archäologie Österreichs 1991;2(2): 11–17.

34. Bagolini B, Dal Ri L, Lippert A, Nothdurfter H. Der Mann im Eis: Die Fundbergung 1992 am Tisenjoch, Gem. Schnals, Südtirol. In: Spindler K, Rastbichler-Zissernig E, Wilfing H, Nedden D, Nothdurfter H, editors. Der Mann im Eis Neue Funde und Ergebnisse. Wien: Springer Verlag; 1995. pp. 3–23.

35. ter Braak CJF, Smilauer P. CANOCO Reference Manual and CanoDraw for Windows User's Guide: Software for Canonical Community Ordination (version 4.5). Ithaca NY, USA: Microcomputer Power; 2002.

36. ter Braak CJF, Prentice IC. A Theory of Gradient Analysis. Adv Ecol Res. 1988;18: 271–317.

37. ter Braak CJF, Wiertz J. On the Statistical Analysis of Vegetation Change: A Wetland Affected by Water Extraction and Soil Acidification. J Veg Sci. 1994;5(3): 361–372.

38. Dickson JH. Bryophytes of the Pleistocene: the British record and its chorological and ecological implications. London: Cambridge University Press; 1973.

39. Dickson JH. Bryophyte Analysis. In: Berglund BE, editor. Handbook of Holocene palaeoecology and palaeohydrology. Chichester: Wiley‐Interscience; John Wiley & Sons Ltd.; 1986. pp. 627–643

40. Miller NG. Tertiary and Quaternary Fossils. In: Schuster RM, editor. New Manual of Bryology: Volume 2. Nichinan: Hattori Botanical Laboratory; 1984. pp. 1194–1232

41. Haeberli W, Kääb A, Wagner S, Mühll DV, Geissler P, Haas JN, et al. Pollen analysis and 14C age of moss remains in a permafrost core recovered from the active rock glacier Murtèl-Corvatsch, Swiss Alps: geomorphological and glaciological implications. Journal of Glaciology. 1999;45(149): 1–8.

42. Krainer K, Bressan D, Dietre B, Haas JN, Hajdas I, Lang K, et al. A 10,300-year-old permafrost core from the active rock glacier Lazaun, southern Ötztal Alps (South Tyrol, northern Italy). Quat Res. 2015;83(2): 324–335.

43. Dickson JH, Mudie PJ, Mackie AP, Coppins B, Hastings R, Hebda RJ, et al. Vascular Plants, Bryophytes, Lichens and Algae from the Kwädąy Dän Ts’ìnchį Discovery Site in Northwestern British Columbia. Kwädąy Dän Ts’ìnchį: Teachings from Long Ago Person Found. Victoria, Canada: Royal British Columbian Museum; 2017. pp. 89–113.

44. Dickson JH, Oeggl K, Stanton D. ‘Forest Moss’: no part of the European Neanderthal diet. Antiquity. 2017;91(359): 1–6.

45. Grims F. Die Laubmoose Österreichs. Wien: Osterreichische Akademis der Wissenschaften; 1999.

46. Akeret Ö, Jacomet S. Analysis of plant macrofossils in goat/sheep faeces from the Neolithic lake shore settlement of Horgen Scheller—an indication of prehistoric transhumance? Veg Hist Archaeobot. 1997;6(4): 235–239.

47. Jacomet S, Leuzinger U, Schibler J. Die Jungsteinzeitliche Seeufersiedlung Arbon Bleiche 3: Umwelt und Wirtschaft. St. Gallen: Department for Education and Culture; Canton of Thurgau; 2004.

48. Ihl C, Barboza PS. Nutritional Value of Moss for Arctic Ruminants: a Test with Muskoxen. J Wildl Manage. 2007;71(3): 752–758.

49. Flø D, Hagvar S. Aerial Dispersal of Invertebrates and Mosses Close to a Receding Alpine Glacier in Southern Norway. Arctic, Antarctic and Alpine Research. 2013;45, 481–490.

50. Spindler K. The man in the ice: the preserved body of a Neolithic man reveals the secrets of the Stone Age. London: Phoenix; 2001.

51. Dierßen K. Distribution, ecological amplitude and phytosociological characterization of European bryophytes. Stuttgart: Schweizerbart; 2001.

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