Abstract
Expanding visitation to Polar regions combined with climate warming increases the potential for alien species introduction and establishment. We quantified vascular plant propagule pressure associated with different groups of travelers to the high-Arctic archipelago of Svalbard, and evaluated the potential of introduced seeds to germinate under the most favorable average Svalbard soil temperature (10°C). We sampled the footwear of 259 travelers arriving by air to Svalbard during the summer of 2008, recording 1,019 seeds: a mean of 3.9 (±0.8) seeds per traveler. Assuming the seed influx is representative for the whole year, we estimate a yearly seed load of around 270,000 by this vector alone. Seeds of 53 species were identified from 17 families, with Poaceae having both highest diversity and number of seeds. Eight of the families identified are among those most invasive worldwide, while the majority of the species identified were non-native to Svalbard. The number of seeds was highest on footwear that had been used in forested and alpine areas in the 3 months prior to traveling to Svalbard, and increased with the amount of soil affixed to footwear. In total, 26% of the collected seeds germinated under simulated Svalbard conditions. Our results demonstrate high propagule transport through aviation to highly visited cold-climate regions and isolated islands is occurring. Alien species establishment is expected to increase with climate change, particularly in high latitude regions, making the need for regional management considerations a priority.
This is a preview of subscription content, access via your institution.


References
Alsos IG, Eidesen PB, Ehrich D, Skrede I, Westergaard K, Jacobsen GH, Landvik J, Taberlet P, Brochmann C (2007) Frequent long-distance plant colonization in the changing Arctic. Science 316:1606–1609
Alsos IG, Gillespie L, Marusik YM (2009) Arctic islands, biology. In: Gillespie R, Clague DA (eds) Encyclopedia of Islands. University of California Press, Berkeley
Alsos IG, Arnesen G, Sandbakk BE, Elven R (2011). http://svalbardflora.net. Accessed 20 Aug 2011
Ashton GV, Riedlecker EI, Ruiz GM (2008) First non-native crustacean established in coastal waters of Alaska. Aquat Biol 3:133–137
Australia and SCAR (Scientific Committee on Antarctic Research) (2007) IPY Aliens in Antarctica. Information paper presented by the Government of Australia and the Scientific Committee on Antarctica Research: information paper 49. Antarctic Treaty Consultative Meeting (ATCM XXX), New Dehli
Barnes DKA, Hodgson DA, Convey P, Allen CS, Clarke A (2006) Incursion and excursion of Antarctic biota: past, present and future. Glob Ecol Biogeogr 15:121–142. doi:10.1111/j.1466-822x.2006.00216.x
Baskin CC, Baskin JM (1998) Seeds: ecology, biogeography, and evolution of dormancy and germination. Academic Press, San Diego
Biosecurity New Zealand (2010) Entering New Zealand. New Zealand Government, New Zealand. http://www.biosecurity.govt.nz/enter. Accessed 6 May 2010
Cappers RTJ, Bekker RM, Jans JEA (2006) Digital seed atlas of the Netherlands. Groningen archaeological studies 4. Barkhuis Publishing, The Netherlands
Carlton JT (1985) Transoceanic and interoceanic dispersal of coastal marine organisms: the biology of ballast water. Mar Biol Ann Rev 23:313–371
Carlson ML, Shephard M (2007) Is the spread of non-native plants in Alaska accelerating? Meeting the challenge: invasive plants in Pacific Northwest ecosystems. U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, Portland, OR
Clifford HT (1956) Seed dispersal on footwear. Proc Bot Soc Br Isles 2:129–131
Colautti RI, Grigorovich IA, MacIsaac HJ (2006) Propagule pressure: a null model for biological invasions. Biol Invasions 8:1023–1037. doi:10.1007/s10530-005-3735-y
Convey P, Frenot Y, Gremmen N, Bergstrom DM (2006) Biological invasions. In: Bergstrom DM, Convey P, Huskies AHL (eds) Trends in Antarctic terrestrial and limnetic ecosystems: Antarctica as a global indicator. Springer, The Netherlands, pp 194–220
Coulson SJ, Refseth D (2004) The terrestrial and freshwater invertebrate fauna of Svalbard (and Jan Mayen). In: Prestrud P, Strøm H, Goldman H (eds) A catalogue of the terrestrial and marine animals of Svalbard. Norsk Polarinstitutt 201, pp 57–122
Curry CH, McCarthy JS, Darragh HM, Wake RA, Todhunter R, Terris J (2002) Could tourist boots act as vectors for disease transmission in Antarctica? J Travel Med 9:190–193
Drake LA, Doblin MA, Dobbs FC (2007) Potential microbial bioinvasions via ships’ ballast water, sediment, and biofilm. Mar Poll Bull 55:333–341
Elvebakk A (2005) A vegetation map of Svalbard on the scale 1:3.5 mill. Phytocoenologia 35:951–967
Elvebakk A, Prestrud P (1996) A catalogue of Svalbard plants, fungi, algae and cyanobacteria. Norsk Polarinstitutt 198, pp 1
Elven R, Elvebakk A (1996) Part 1. Vascular plants. In Elvebakk A, Prestrud P (eds) A catalogue of Svalbard plants, fungi, algae, and cyanobacteria. Norsk Polarinstitutt 198, pp 9–55
Elven R, Murray DF, Razzhivin V, Yurtsev BA (2011) Checklist of the panarctic flora (PAF). CAFF/University of Oslo, Oslo. http://gbif.no/paf. Accessed 20 Aug 2011
Falinski JB (1972) Anthropochory in xerothermic grasslands in the light of experimental data. Acta Soc Bot Pol 41:357–368
Forbes BC, Monz C, Tolvanen A (2004) Ecological impacts of tourism in terrestrial polar ecosystems. In: Buckley R (ed) Environmental impacts of ecotourism. CAB International, Oxfordshire, pp 155–170
Fredga K, Jaarola M, Ims RA, Steen H, Yoccoz NG (1990) The ‘common vole’ in Svalbard identified as Microtus epiroticus by chromosome analysis. Polar Res 8:283–290
Frenot Y, Chown SL, Whinam J, Selkirk PM, Convey P, Skotnicki M, Bergstrom DM (2005) Biological invasions in the Antarctic: extent, impacts and implications. Biol Rev 80:45–72. doi:10.1017/s1464793104006542
Gillespie GD (1985) Hybridization, introgression, and morphometric differentiation between Mallard (Anas platyrhynchos) and Grey Duck (Anas superciliosa) in Otago, New Zealand. The Auk 102:459–469
Gollasch S (2002) The importance of ship hull fouling as a vector of species introductions in to the North Sea. Biofouling 18:105–121
Governor of Svalbard (2006) Report on tourism statistics from Svalbard 1997–2006. Svalbard Sysselmannen. http://www.sysselmannen.no/hovedEnkel.aspx?m=45275. Accessed 10 Mar 2009
Hanssen-Bauer I (2002) Temperature and precipitation in Svalbard 1912–2050: measurements and scenarios. Polar Rec 38:225–232
Higashino PK, Guyer W, Stone CP (1983) The Kilauea wilderness marathon and crater rim runs: sole searching experiences. Newsl Hawaii Bot Soc 22:22–28
Hines AH, Ruiz GM (2000) Biological invasions at cold-water coastal ecosystems: ballast- mediated introductions in Port Valdez/Prince William Sound. Final Report to Regional Citizens Advisory Council of Prince William Sound
Hodkinson DJ, Thompson K (1997) Plant dispersal: the role of man. J Appl Ecol 34:1484–1496
Hughes K, Lee J, Ware C, Kiefer K, Bergstrom D (2010) Impact of anthropogenic transportation to Antarctica on alien seed viability. Polar Biol 33:1125–1130. doi:10.1007/s00300-010-0801-4
IAATO (International Association of Antarctica Tour Operators) (2010) Boot, clothing and equipment decontamination guidelines for small boat operators. IAATO. http://www.iaato.org/guidelines.html. Accessed 30 May 2010
ISSG (Invasive Species Specilist Group) (2010) Global invasive species database. http://www.issg.org/database/welcome/disclaimer.asp. Accessed 3 June 2010
Jónsdóttir IS (2005) Terrestrial ecosystems on Svalbard: heterogeneity, complexity and fragility from an arctic island perspective. Proc Roy Ir Acad 105B:155–165
Kaltenborn BP (2000) Arctic-alpine environments and tourism: can sustainability be planned? lessons learned on Svalbard. Mt Res Dev 20:28–31
Kirkbride JH Jr, Gunn CR, Dallwitz MJ (2006) Family guide for fruits and seeds. United States Department of Agriculture, Washington, DC. http://nt.ars-grin.gov/sbmlweb/OnlineResources/frsdfam/Index.cfm. Accessed 12 May 2010
Lee JE, Chown SL (2009a) Breaching the dispersal barrier to invasion: quantification and management. Ecol Appl 19:1944–1959. doi:10.1890/08-2157.1
Lee JE, Chown SL (2009b) Quantifying the propagule load associated with the construction of an Antarctic research station. Antarct Sci 21:471–475. doi:10.1017/S0954102009990162
Liška J, Soldán Z (2004) Alien vascular plants recorded from the barentsburg and pyramiden settlements, Svalbard. Preslia 76:279–290
Lockwood JL, Cassey P, Blackburn T (2005) The role of propagule pressure in explaining species invasions. Trends Ecol Evol 20:223–228
Lockwood JL, Hoopes MF, Marchetti MP (2007) Invasion ecology. Blackwell, Malden, USA, pp 59–76
McCullough DG, Work TT, Cavey JF, Liebhold AM, Marshall D (2006) Interceptions at nonindigenous plant pests at US ports of entry and border crossings over a 17-year period. Biol Invasions 8:611–630
McNeill M, Phillips C, Young S, Shah F, Aalders L, Bell N, Gerard E, Littlejohn R (2011) Transportation of nonindigenous species via soil on international aircraft passengers’ footwear. Biol Invasions. doi:10.1007/s10530-011-9964-3
Milbau A, Graae BJ, Shevtsova A, Nijs I (2009) Effects of a warmer climate on seed germination in the subarctic. Ann Bot 104:287–296
Müller E, Cooper EJ, Also IG (2011) Germinability of arctic plants is high in perceived optimal conditions but low in the field. Botany 89:337–348
Pickering C, Mount A (2010) Do tourists disperse weed seed? a global review of unintentional human-mediated terrestrial seed dispersal on clothing, vehicles and horses. J Sustain Tour 18:239–256. doi:10.1080/09669580903406613
Powell RH (1968) Harmful plant species entering New Zealand 1963–1967. NZ J Bot 6:396–401
Puth LM, Post DM (2005) Studying invasion: have we missed the boat? Ecol Lett 8:715–721
Pyšek P (1998) Is there a taxonomic pattern to plant invasions? Oikos 82:282–294
R Development Core Team (2008) R 2.7.2. A language and environment. Vienna, Austria. Available from http://cran.r-project.org
Rowntree JK, Cowan RS, Leggett M, Ramsay MM, Fay MF (2010) Which moss is which? Identification of the threatened moss Orthodontium gracile using molecular and morphological techniques. Conserv Genet 11:1033–1042
SCAR (Scientific Committee on Antarctic Research) (2010) Preliminary results from the International Polar Year Program: Aliens in Antarctica. Working paper 4 presented to the Committee on Environmental Protection. Antarctic Treaty Consultative Meeting XXXIII, Punta del Este, Uruguay
Scherrer D, Körner C (2010) Infra-red thermometry of alpine landscapes challenges climatic warming projections. Global Change Biol 16:2602–2613
Schmidt W (1989) Plant dispersal by motor cars. Vegetatio 80:147–152
Shevtsova A, Graae BJ, Jochum T, Milbau A, Kockelbergh F, Beyens L, Nijs I (2009) Critical periods for impact of climate warming on early seedling establishment in subarctic tundra. Global Change Biol 15:2662–2680
Sørensen T (1941) Temperature relations and phenology of the northeast greenland flowering plants. Commission for Scientific Research in Greenland, Copenhagen
Statistics Greenland (2009) Flight passenger statistics. Department of Finances, Greenland. http://www.stat.gl/Statistik/Turisme/tabid/99/language/en-US/Default.aspx. Accessed 19 Jan 2010
Thompson K, Bakker JP, Bekker RM (1997) The soil seed banks of north west Europe: methodology, density and longevity. Cambridge University Press, Cambridge
Trudgill DL, Squire GR, Thompson K (2000) A thermal time basis for comparing the germination requirements of some British herbaceous plants. New Phytol 145:107–114
Walker DA, Raynolds MK, Daniels FJA, Einarsson E, Elvebakk A, Gould WA, Katenin AE, Kholod SS, Markon CJ, Melnikov ES, Moskalenko NG, Talbot SS, Yurtsev BA (2005) The circumpolar arctic vegetation map. J Veg Sci 16:267–282
Whinam J, Chilcott N, Bergstrom DM (2005) Subantarctic hitchhikers: expeditioners as vectors for the introduction of alien organisms. Biol Conserv 121:207–219
Wichmann MC, Alexander MJ, Soons MB, Galsworthy S, Dunne L, Gould R, Fairfax C, Niggemann M, Hails RS, Bullock JM (2009) Human-mediated dispersal of seeds over long distances. Proc Roy Soc Lond B 276:523–532. doi:10.1098/rspb.2008.1131
Williamson M (1996) Biol invasions. Chapman & Hall, London, pp 28–52
Acknowledgments
We thank the many people who assisted in footwear sampling at the Svalbard airport, and the 259 participants who kindly gave up their shoes for cleaning after arrival. We would also like to thank the staff at the Svalbard Airport for allowing us to conduct the study on airport premises. Financial assistance for this project was received from the Svalbard Environmental Protection Fund. This project is closely aligned with the IPY Aliens in Antarctica Project. Finally, we wish to extend our thanks to three anonymous reviewers for their comments on the manuscript.
Author information
Authors and Affiliations
Corresponding author
Rights and permissions
About this article
Cite this article
Ware, C., Bergstrom, D.M., Müller, E. et al. Humans introduce viable seeds to the Arctic on footwear. Biol Invasions 14, 567–577 (2012). https://doi.org/10.1007/s10530-011-0098-4
Received:
Accepted:
Published:
Issue Date:
DOI: https://doi.org/10.1007/s10530-011-0098-4
Keywords
- Alien
- Non-indigenous
- Dispersal
- Germination
- Human-mediated dispersal
- Propagule pressure