Biological Invasions

, Volume 15, Issue 9, pp 1933–1946 | Cite as

Telling a different story: a global assessment of bryophyte invasions

  • Franz Essl
  • Klaus Steinbauer
  • Stefan Dullinger
  • Thomas Mang
  • Dietmar Moser
Original Paper


We assess and review spatio-temporal patterns, habitat affiliations, pathways, impacts, and management experience of bryophyte invasions in extra-tropical countries and regions (n = 82) from five continents and maritime islands spanning both hemispheres. Distribution data were extracted and critically checked from a wide range of sources and supplemented with data on biology and introduction history. We identified 139 bryophytes species which we consider to be alien in at least one of our study regions (106 mosses, 28 hepatics and 5 hornworts). Numbers of average alien bryophyte species are significantly higher on islands than in continental regions of similar size, and peak on maritime islands. Cumulative numbers of first records have grown slowly until 1950 and have strongly increased since then. Accidental import as hitch-hiker (34 species) or with ornamental plants (27 species) constitute the most important introduction pathways. We found a remarkably high contribution from distant donor regions to alien bryophyte floras, especially from the complementary hemisphere. Most alien bryophytes prefer strongly modified habitats (e.g. ruderal vegetation, roadsides, lawns), and only few natural ecosystems (forests, rocks) are regularly invaded. Evidence for an ecological impact of bryophyte invasions is scarce and competitive replacement of native moss species, or vascular plant seedlings, by alien bryophytes has only been documented in a few cases. We conclude that bryophytes differ profoundly in many respects from vascular plants, and so do their invasion patterns at large scale. Our global bryophyte invasion state assessment provides the basis for future, more explicit considerations of this largely neglected taxonomic group in invasion ecology, a step we suggest to be urgently needed as studying them might provide novel insights into patterns and processes of plant invasions in general.


Alien species Bryophyta Ecosystem Hemisphere Introduction Invasion Naturalization Pathway Region of origin 



Many colleagues have contributed their knowledge to the underlying data set (in brackets: region for which data have been provided): J. Beever (Chathams, New Zealand), L. Cave (Tasmania), H. Deguchi (Japan), A. Fife (Chathams, New Zealand), J.-P. Frahm (Germany, Macaronesia), D. Glenny (New Zealand), J. M. González-Mancebo (Canary Islands), M. Hill (UK), J. Klinck (Denmark), P. Lambdon (Ascension, St. Helena), J. Larrain (Chile), N. Miller (USA), F. Müller (Chile, Argentina), R. Porley (UK), R. A. Pursell (USA), R. Seppelt (Australia), M. Sabovljevic (Serbia), J. Shevock (California), J. S. Song (South Korea, Japan), A. Stebel (Poland), J.-D. Yang (Taiwan), R. Zander (USA). Their contributions have been extremely helpful. The comments of two anonymous reviewers are highly appreciated.

Supplementary material

10530_2013_422_MOESM1_ESM.doc (198 kb)
Supplementary material 1 (DOC 198 kb)
10530_2013_422_MOESM2_ESM.csv (43 kb)
Appendix S2: Alien bryophyte species, their invasion status (nat = naturalized, cas = casual, crypt = cryptogenic), year of first record (FR, if known), native range (continents), pathways, habitats colonized, and substrate type colonized in the 82 regions included in this study. Supplementary material 2 (CSV 43 kb)
10530_2013_422_MOESM3_ESM.doc (181 kb)
Supplementary material 3 (DOC 181 kb)
10530_2013_422_MOESM4_ESM.doc (63 kb)
Supplementary material 4 (DOC 63 kb)


  1. Aleffi M, Ricci S, Tacchi R (2010) Hypopterygium tamarisci (Sw.) Brid. ex Müll. Hal. (Hypopterygiaceae, Bryopsida), new to Italy. Cryptogam Bryol 31:293–295Google Scholar
  2. Arts T (2001) A revision of the Splachnobryaceae (Musci). Lindbergia 26:77–96Google Scholar
  3. Australian Biological Resource Study (ed) (2006) Flora of Australia, volume 51: mosses. CSIRO Publishing, CollingwoodGoogle Scholar
  4. Australian National Botanic Garden (2012) Bryophytes. Accessed 20 Feb 2012
  5. Barry RG, Chorley RJ (1992) Atmosphere, weather, and climate. Routledge, LondonGoogle Scholar
  6. Beever J, Allison KW, Child J (1992) The mosses of New Zealand. The University of Otago Press, DunedinGoogle Scholar
  7. BFNA Editorial Committee (2011) Bryophyte flora of North America. Accessed 11 Nov 2011
  8. Biermann R, Daniels FJA (1997) Changes in a lichen-rich dry sand grassland vegetation with special reference to lichen synusiae and Campylopus introflexus. Phytocoenologia 27:257–273Google Scholar
  9. Bobbink R, Hicks K, Galloway J, Spranger T, Alkemade R, Ashmore M, Bustamante M, Cinderby S, Davidson E, Dentener F, Emmett B, Erisman JW, Fenn M, Gilliam F, Nordin A, Pardo L, De Vries W (2010) Global assessment of nitrogen deposition effects on plant terrestrial biodiversity: a synthesis. Ecol Appl 20:30–59PubMedCrossRefGoogle Scholar
  10. Chytrý M, Pyšek P, Tichy L, Knollova I, Danihelka J (2005) Invasions by alien plants in the Czech Republic: a quantitative assessment across habitats. Preslia 77:339–354Google Scholar
  11. CIA (2009) The World Factbook 2009. Central Intelligence Agency, Washington, DCGoogle Scholar
  12. Colpa H, van Zanten B (2009) Scopelophila cataractae op stortplaats voormalig kamp Westerbork [Scopelophila cataractae discovered on site of 60 year old dump of zinc–carbon batteries]. Buxbaumiella 84:1–6Google Scholar
  13. Cronk QCB, Fuller JL (1995) Plant invaders. Chapman & Hall, LondonGoogle Scholar
  14. DAISIE (2012) Delivering alien invasive species inventories for Europe. Accessed 12 Feb 2012
  15. Denslow JS, Space JC, Thomas PA (2009) Invasive exotic plants in the tropical pacific islands: patterns of diversity. Biotropica 41:162–170CrossRefGoogle Scholar
  16. Dickson JH (1967) Pseudoscleropodium purum (Limpr.) Fleisch. on St. Helena and its arrival on Tristan da Cunha. Bryologist 70:267–268Google Scholar
  17. Dobson AT (1975) Sphagnum subnitens, S. squarrosum, and Drepanocladus revolvens in New Zealand mires. NZ J Bot 13:169–171CrossRefGoogle Scholar
  18. Ellis EC, Antill EC and Kreft H (2012) All is not loss: plant biodiversity in the Anthropocene. PLoS ONE 7. doi: 10.1371/journal.pone.0030535
  19. Equiha M, Usher MB (1993) Impacts of carpets of the invasive moss Campylopus introflexus on Calluna vulgaris regeneration. J Ecol 81:359–365CrossRefGoogle Scholar
  20. Essl F, Lambdon P (2009) The alien bryophytes and lichens of Europe. In: DAISIE (ed). Springer, Berlin, pp 29–42Google Scholar
  21. Essl F, Lambdon P, Rabitsch W (2011) Bryophytes and lichens. In: Simberloff D, Rejmanek M (eds). University of California Press, Berkeley, pp 81–85Google Scholar
  22. Faraway JJ (2006) Extending the linear model with R: generalized linear, mixed effects and nonparametric regression models. Chapman & Hall, Boca RatonGoogle Scholar
  23. Frahm JP (1973) Über Vorkommen und Verbreitung von Lunularia cruciata (L.) Dum. in Deutschland. Herzogia 2:396–409Google Scholar
  24. Frahm JP, Klaus D (2001) Bryophytes as indicators of recent climate fluctuations in Central Europe. Lindbergia 26:97–104Google Scholar
  25. Frey W, Frahm JP, Fischer E, Lobin W (2006) The liverworts, mosses and ferns of Europe. Harley Books, ColchesterGoogle Scholar
  26. Gradstein SR, Vana J (1987) On the occurrence of Laurasian liverworts in the tropics. Mem NY Bot Gard 45:388–425Google Scholar
  27. Hall J, Fukuda S, Hoe WJ (1995) Riccia fluitans (Hepaticae, Ricciaceae): first observation in Hawaii. Fragm Florist Geobot 40:233–234Google Scholar
  28. Hasse T (2007) Campylopus introflexus invasion in a dune grassland: succession, disturbance and relevance of existing plant invader concepts. Herzogia 20:305–315Google Scholar
  29. Hassel K, Söderström L (2005) The expansion of the alien mosses Orthodontium lineare and Campylopus introflexus in Britain and continental Europe. J Hattori Bot Lab 97:183–193Google Scholar
  30. Hedenäs L, Herben T, Rydin H, Söderström L (1989) Ecology of the invading moss Orthodontium lineare in Sweden: spatial distribution and population structure. Holarctic Ecol 12:163–172Google Scholar
  31. Herben T (1994) Local rate of spreading and patch dynamics of an invasive moss species, Orthodontium lineare. J Bryol 18:115–125Google Scholar
  32. Hill MO, Baker R, Broad G, Chandler PJ, Copp GH, Ellis J, Jones D, Hoyland C, Laing I, Longshaw M, Moore N, Parrott D, Pearman D, Preston C, Smith RM, Waters R (2005) Audit of non-native species in England. English Nature research reportsGoogle Scholar
  33. Hill MO, Bell N, Bruggeman-Nannenga MA, Brugués M, Cano MJ, Enroth J, Flatberg KI, Frahm JP, Gallego MT, Garilleti R, Guerra J, Hedenäs L, Holyoak DT, Hyvönen J, Ignatov MS, Lara F, Mazimpaka V, Muñoz J, Söderström L (2006) An annotated checklist of the mosses of Europe and Macaronesia. J Bryol 28:198–267CrossRefGoogle Scholar
  34. Holyoak D, Lockhart N (2009) Australasian bryophytes introduced to South Kerry with tree ferns. Field Bryol 98:3–7Google Scholar
  35. Hulme PE, Bacher S, Kenis M, Klotz S, Kühn I, Minchin D, Nentwig W, Olenin S, Panov V, Pergl J, Pyšek P, Roques A, Sol D, Solarz W, Vilà M (2008) Grasping at the routes of biological invasions: a framework for integrating pathways into policy. J Appl Ecol 45:303–341CrossRefGoogle Scholar
  36. Ketner-Oostra R, Sýkora KV (2000) Vegetation succession and lichen diversity on dry coastal calcium-poor dunes and the impact of management experiments. J Coast Conserv 6:191–206CrossRefGoogle Scholar
  37. Ketner-Oostra R, Sýkora KV (2004) Decline of lichen-diversity in calcium poor coastal dune vegetation since the 1970s, related to grass and moss encroachment. Phytocoenologia 34:521–549CrossRefGoogle Scholar
  38. Ketner-Oostra R, Sýkora KV (2008) Vegetation in a lichen-rich inland drift sand area in the Netherlands. Phytocoenologia 38:267–286CrossRefGoogle Scholar
  39. Klinck J (2009) The alien invasive species Campylopus introflexus in the Danish coastal dune system. Master thesis, Department Biology, Section for Ecology and Evolution, Copenhagen UniversityGoogle Scholar
  40. Kucera J (1999) Didymodon australasiae var. umbrosus in the Czech Republic, with a review of recent records from Central Europe. J Bryol 21:71–77Google Scholar
  41. Kueffer C, Daehler CC, Torres-Santana CW, Lavergne C, Meyer JY, Otto R, Silva L (2010) A global comparison of plant invasions on oceanic islands. Perspect Plant Ecol Evol Syst 12:145–161CrossRefGoogle Scholar
  42. Lambdon PW, Pyšek P, Basnou C, Hejda M, Arianoutsou M, Essl F, Jarošik V, Pergl J, Winter M, Anastasiu P, Andriopoulos P, Bazos I, Brundu G, Celesti-Grapow L, Chassot P, Delipetrou P, Josefsson M, Kark S, Klotz S, Kokkoris Y, Kühn I, Marchante H, Perglova I, Pino J, Vilá M, Zikos A, Roy D, Hulme PE (2008) Alien flora of Europe: species diversity, temporal trends, geographical patterns and research needs. Preslia 80:101–149Google Scholar
  43. Landcare Research (2012) A new moss flora of New Zealand. Accessed 19 March 2012
  44. Magill RE (2010) Moss diversity: new look on old numbers. Phytotaxa 9:167–174Google Scholar
  45. Magill RE, van Rooy J (1998) Flora of Southern Africa—Bryophyta—part 1, Musci. National Botanical Institute, PretoriaGoogle Scholar
  46. Matteri CM (2003) Los musgos (Bryophyta) de Argentina. Trop Bryol 24:33–100Google Scholar
  47. Mikulášková E, Fajmonová Z, Hájek M (2012) Invasion of the moss Campylopus introflexus into central European habitats. Preslia 84:863–886Google Scholar
  48. Miller HA (1967) Oddments of Hawaiian bryology. J Hattori Bot Lab 30:271–276Google Scholar
  49. Miller NG (2009) Mosses adventive and naturalized in the northeastern United States: new examples and new distributional records. Rhodora 111:218–230CrossRefGoogle Scholar
  50. Miller NG, Robinson SC (2010) Introduction and recent range expansion in the moss Ptychomitrium serratum (Ptychomitriaceae) in the southern and eastern United States. Botany 88:336–344CrossRefGoogle Scholar
  51. Miller NG, Trigoboff N (2001) A European feather moss, Pseudoscleropodium purum, naturalized widely in New York State in cemeteries. Bryologist 10:98–103CrossRefGoogle Scholar
  52. Müller F (2002) Ein Freilandnachweis von Didymodon australasiae var. umbrosus in Deutschland. Herzogia 15:187–190Google Scholar
  53. Mutke J, Geffert JL (2010) Keep on working: the uneven documentation of regional moss floras. Trop Bryol 31:7–13Google Scholar
  54. Noguchi A (1987–1994) Illustrated moss flora of Japan, part 1–5. The Hattori Botanical Laboratory, HiroshimaGoogle Scholar
  55. Norris DH, Koponen T, Piippo S (1999) Bryophyte flora of the Huon Penninsula, Papua New Guinea, LXVI. Meesiaceae (Musci), with lists of boreal to temperate disjunct, bipolar, and widely distributed species. Ann Bot Fenn 36:257–263Google Scholar
  56. Paton JA (1999) The liverwort flora of the British Isles. Harley Books, ColchesterGoogle Scholar
  57. Pemberton RW, Liu H (2009) Marketing time predicts naturalization of horticultural plants. Ecology 90:69–80PubMedCrossRefGoogle Scholar
  58. Porley R, Haynes T (2009) An update on the alien liverwort Lophocoela semiteres (Lehm.) Mitt. and its spread in Britain and Ireland. Field Bryol 99:3–9Google Scholar
  59. Porley R, Hodgetts N (2005) Mosses and liverworts. New naturalist series. Harper Collins, LondonGoogle Scholar
  60. Pyšek P, Richardson DM, Rejmánek M, Webster GL, Williamson M, Kirschner J (2004) Alien plants in checklists and floras: towards better communication between taxonomists and ecologists. Taxon 53:131–143CrossRefGoogle Scholar
  61. Pyšek P, Richardson DM, Pergl J, Jarošik V, Sixtová Z, Weber E (2008) Geographical and taxonomical biases in invasion ecology. Trends Ecol Evol 23:237–244PubMedCrossRefGoogle Scholar
  62. Pyšek P, Jarošík V, Hulme PE, Kühn I, Wild J, Arianoutsou M, Bacher S, Chiron F, Didžiulis V, Essl F, Genovesi P, Gherardi F, Hejda M, Kark S, Lambdon PW, Desprez-Loustau AM, Nentwig W, Pergl J, Poboljšaj K, Rabitsch W, Roques A, Roy DB, Solarz W, Vilà M, Winter M (2010) Disentangling the role of environmental and human pressures on biological invasions. Proc Natl Acad Sci USA 107:12157–12162PubMedCrossRefGoogle Scholar
  63. Reaser JK, Meyerson LA, Cronk Q, De Poorter M, Eldrege LG, Green E, Kairo M, Latasi P, Mack RN, Mauremootoo J, O’Dowd D, Orapa W, Sastroutomo S, Saunders A, Shine C, Thrainsson S, Vaiutu L (2007) Ecological and socioeconomic impacts of invasive alien species in island ecosystems. Environ Conserv 34:98–111CrossRefGoogle Scholar
  64. Richards PW (1963) Campylopus introflexus (Hedw.) Brid. and C. polytrichoides De Not. in the British Isles; a preliminary account. Trans Br Bryol Soc 4:404–417CrossRefGoogle Scholar
  65. Rumsey FJ (1992) The status of Tortula freibergii in the British Isles. J Bryol 17:371–373Google Scholar
  66. Schirmel J (2011) Response of the grasshopper Myrmeleotettix maculatus (Orthoptera: Acrididae) to invasion by the exotic moss Campylopus introflexus in acidic coastal dunes. J Coast Conserv 15:159–162CrossRefGoogle Scholar
  67. Schirmel J, Timler L, Buchholz S (2011) Impact of the invasive moss Campylopus introflexus on carabid beetles (Cleoptera: Carabidae) and spiders (Araneae) in acidic coastal dunes at the southern Baltic Sea. Biol Invasions 13:605–620CrossRefGoogle Scholar
  68. Schofield WB (1997) Bryophytes unintentionally introduced to British Columbia. Botanical Electronic News 162.
  69. Seppelt RD, Cave LH (2011) Introduced mosses in the flora of Tasmania, II. Kindbergia praelonga (Bryopysida: Brachytheciaceae). Kannunnah 4:82–88Google Scholar
  70. Seppelt RD, Cave LH, Carter BE (2011) Introduced mosses in the flora of Tasmania I. Scleropodium and Pseudoscleropodium (Bryopsida: Brachytheciaceae). Kannunnah 4:72–81Google Scholar
  71. Smith AJE (2004) The moss flora of the British Isles. Cambridge University Press, CambridgeGoogle Scholar
  72. Söderström L (1992) Invasions and range expansions and contractions of bryophytes. In: Bates JW, Farmer AM (eds). Clarendon Press, Oxford, pp. 131–158Google Scholar
  73. Söderström L, Urmi E, Váňa J (2002) Distribution of Hepaticae and Anthocerotae in Europe and Macaronesia. Lindbergia 27:3–47Google Scholar
  74. Stieperaere H, Jacques A (1995) The spread of Orthodontium lineare and Campylopus introflexus in Belgium. Belg J Bot 128:117–123Google Scholar
  75. R Development Core Team (2012) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna.
  76. Vilà M, Espinar J, Hejda M, Hulme P, Jarošík V, Maron J, Pergl J, Schaffner U, Sun Y, Pyšek P (2011) Ecological impacts of invasive alien plants: a meta-analysis of their effects on species, communities and ecosystems. Ecol Lett 14:702–708PubMedCrossRefGoogle Scholar
  77. Vogels J, Nijssen M, Verberk W, Esselink H (2005) Effects of moss encroachment by Campylopus introflexus on soil-entomofauna of dry-dune grasslands (Ciolo-Corynephoretum). Proc Neth Entomol Soc 16:71–80Google Scholar
  78. Waite M (2007) Mosses of Hawaii Volcanoes National Park. Technical report 153, Pacific Cooperative Studies Unit, University of HawaiiGoogle Scholar
  79. Weber E (2003) Invasive plant species of the world. CABI Publishing, WallingfordGoogle Scholar

Copyright information

© Springer Science+Business Media Dordrecht 2013

Authors and Affiliations

  • Franz Essl
    • 1
  • Klaus Steinbauer
    • 1
  • Stefan Dullinger
    • 2
    • 3
  • Thomas Mang
    • 2
    • 3
  • Dietmar Moser
    • 1
    • 3
  1. 1.Environment Agency AustriaViennaAustria
  2. 2.Department of Conservation Biology, Vegetation and Landscape Ecology, Faculty Centre of BiodiversityUniversity of ViennaViennaAustria
  3. 3.Vienna Institute for Nature Conservation and AnalysesViennaAustria

Personalised recommendations