1 Supplementary Material 2 3 This material is available as part of the online article from: 4 Ozinga et al. Dispersal failure contributes to plant losses in NW Europe 5 http://www.blackwell-synergy.com/doi/full/10.1111/j.1461-0248.XXXX.XXXXX.x 6 7 Appendix S1 Possible confounding effects 8 Appendix S2 Historical overview of changes in dispersal infrastructure in the landscape 9 of Northwest Europe. 10 11 Please note: Blackwell Publishing is not responsible for the content or functionality of 12 any supplementary materials supplied by the authors. Any queries (other than missing 13 material) should be directed to the corresponding author for the article. 14 15 1 1 2 Appendix S2: Historical overview of changes in dispersal 3 infrastructure in the landscape of Northwest Europe 4 5 Several vectors transport seeds between sites, including water, wind, birds and 6 large mammals, each with their own characteristics (Hodgson & Grime 1990; Fischer et 7 al. 1996; Poschlod & Bonn 1998; Boedeltje et al. 2003; Manzano & Malo 2006; Nathan 8 2006). At the landscape level, these dispersal vectors act like a complex ‘dispersal 9 infrastructure’. In Northwest Europe, wind patterns and bird migrations remained almost 10 unchanged throughout the 19th and 20th centuries (KNMI 2005 respectively LWVT / 11 SOVON 2002), while the exchange of large mammals and water between sites has 12 greatly decreased. These changes mostly took place some 50 to 150 years ago (Ridley 13 1930; Beaufoy et al. 1994; Dynesius & Nilsson 1994; European Environment Agency 14 2003; Poschlod et al. 2005; see Poschlod & Bonn 1998 for a comprehensive review). 15 16 Free-ranging or herded large mammals 17 For many plant species, the retention time of seeds by large mammals (both 18 externally in the fur and internally in the digestive tract) is long enough to enable seeds to 19 be transported across corridors over distances of many kilometres (Fischer et al. 1996; 20 Pakeman 2001; Myers et al. 2004; Manzano & Malo 2006). Of the large herbivores that 21 roamed the plains and forests of Pleistocene NW Europe, most large mammalian species 22 and all ‘megaherbivores’ became extinct (Anderson 1984; Stuart 1991, 2005). Since there 2 1 is a positive log-linear relation between body weight and dispersal distances (as regards 2 both median and maximum dispersal distances, Sutherland et al. 2000), the most effective 3 long-distance dispersal vectors among the mammals have thereby probably disappeared. 4 Only four widely distributed ungulate species have remained (in order of increasing 5 median dispersal distances): Roe deer (Capreolus capreolus), Fallow deer (Dama dama), 6 Wild boar (Sus scrofa) and Red deer (Cervus elaphus). The natural migration of these 7 remaining wild mammalian species is currently severely hampered in most European 8 landscapes (Wallis de Vries 1995; Groot Bruinderink et al. 2003). 9 Grazing by livestock, i.e. horses, cattle, sheep, goats and pigs, can be regarded as 10 a modern analogy of seed dispersal by the original fauna (Janzen & Martin 1982, Olff & 11 Ritchie 1998). Until the beginning of the 20th century, it was common for farmers to herd 12 their livestock on a daily basis on the unfenced pastures, covering distances of 13 approximately 0.5 to 10 km per day. The highly branched networks of drift-roads for 14 livestock around the villages remained in use for many centuries (Slichter Van Bath 15 1980; Jager 1985; Bielemans 1987; Hillegers 1993; Spek 2004). 16 It was also common to transport livestock between regions (over distances of 20- 17 200 km) for trade (Wiese 1966; Bieleman 1987; Hillegers 1993; Gijsbers 1999). These 18 migratory livestock systems (transhumance) existed in large parts of Europe (Ruiz & 19 Ruiz 1986; Whittaker 1988; Poschlod & Bonn 1998; Bruun & Fritbøger 2002; Bunce et 20 al. 2004). Although the seasonal transhumance was potentially important for the 21 migration of plant species over very long distances (>100 km; Fischer et al. 1996; 22 Poschlod & Bonn 1998; Manzano & Malo 2006), the herded livestock within regions was 23 more important for seed dispersal in a quantitative sense. Interregional drift of livestock 3 1 was restricted to a few main drove-roads. By contrast, the daily, local herding of livestock 2 encompassed much larger areas with a higher frequency (Jager 1985; Gijsbers 1999; 3 Spek 2004). 4 Nowadays, free-ranging or herded livestock grazing has become rare in 5 Northwest-Europe. In the Netherlands its spatial coverage has been reduced by more than 6 90% (Slichter Van Bath 1980; Bielemans 1987; Spek 2004) and by 75% in many other 7 parts of Northwest Europe (Beaufoy et al. 1994; Bignal & McCracken 1996; Poschlod & 8 Bonn 1998; Bruun & Fritbøger 2002; Bunce et al. 2004). Free-ranging or herded 9 livestock grazing has been replaced by grazing in fenced fields or livestock housing. This 10 in turn has greatly reduced the potential for mammal-assisted dispersal of plants between 11 sites (Fischer et al. 1996; Poschlod & Bonn 1998; Manzano & Malo 2006). 12 13 14 Free-running and inundating water Palaeo-ecological evidence shows that prior to human inference, the hydrology of 15 many European lowland rivers was dynamic, with multi-braided channels influencing 16 large stretches of land (Brown 2002). In many floodplains in Northwest Europe, 17 inundations were tolerated, and from the Late Middle Ages onwards even stimulated due 18 to the positive effects on grassland productivity (Rackham 1986; Ellenberg 1988; Pott 19 1995; Konold 1997; Poschlod & Bonn 1998). In rivers and brooks throughout Europe, 20 however, natural flood regimes have been altered severely by large flow- and flood- 21 control projects implemented during the 19th and 20th century (Dynesius & Nilsson 22 1994; Lytle & Poff 2004; Nilsson et al. 2005). As a result, the area affected by frequent 23 inundations has been greatly decreased over the last two centuries, in the Netherlands 4 1 (Gottschalk 1977; Kalweit 1993) as well as in many other parts of Europe (Dynesius & 2 Nilsson 1994; Brown 2002; Lytle & Poff 2004; Nilsson et al. 2005). It is estimated that 3 more than 90% of European floodplains are now ‘cultivated’ and therefore functionally 4 ‘extinct’ (Tockner & Stanford 2002). For example, it has been estimated for parts of the 5 Netherlands that until 1860, up to 60% of the area was inundated periodically with 6 surface- or groundwater during the winter period, while this area is now reduced to less 7 than 1% (Corporaal et al. 2002). The restriction of transversal and longitudinal water 8 flows has therefore greatly reduced the potential of seed dispersal between sites 9 (Dynesius & Nilsson 1994; Poschlod & Bonn 1998; Jansson et al. 2000; Lytle & Poff 10 2004; Boedeltje et al. 2003). 11 12 13 References 14 Anderson, E. (1984). Who's who in the Pleistocene: A mammalian bestiary. In: 15 Quaternary extinctions, a prehistoric revolution (eds. Martin, P.S. & R.G. Klein). 16 Univ. of Arizona Press, Tucson, pp. 40-89. 17 Beaufoy, G., Baldock, D. & Clark, J. (1994). The nature of farming. Low intensity 18 farming systems in nine European countries. Joint Nature Conservation 19 Committee, London. 20 21 22 23 Bieleman, J. (1987). Agrarian change in the Dutch province of Drenthe 1600-1910. PhDthesis University of Wageningen. [in Dutch with English summary]. Bignal, E.M. & McCracken, D.I. (1996). Low-intensity farming systems in the conservation of the countryside. J. Appl. Ecol. 33, 413-424. 5 1 Boedeltje, G., Bakker, J.P., Bekker, R.M., Van Groenendael J.M. & Soesbergen, M. 2 (2003). Plant dispersal in a lowland stream in relation to occurrence and three 3 specific life-history traits of species in the species pool. J. Ecol. 91, 855-866. 4 5 6 7 Brown, A.G. (2002). Learning from the past: palaeohydrology and palaeoecology. Freshwat. Biol. 47, 817-829. Bruun, H.H. & Fritzbøger, B. (2002). The past impact of livestock husbandry on dispersal of plant seeds in the landscape of Denmark. Ambio 31, 425-431. 8 Bunce, R.G.H., Pérez-Soba, M., Jongman, R.H.G., Gómez Sal, A., Herzog, F. & Austad, 9 I., Eds. (2004). Transhumance and biodiversity in European mountains. Report of 10 the EU-FP5 project Transhumount (EVK2-CT-2002-80017). IALE Publication 11 Series 1, 1-321. 12 Corporaal, A., Geerlink, H, Huyskes, H., Waaijenberg, J. & de Groot, G. (2002). 13 Spongiteit, een kwestie van volhouden; Vasthouden is beter dan tegenhouden: een 14 verkenning. Alterra/DLG-rapport 422, Wageningen. [Hydrological resilience in 15 Dutch river catchments; in Dutch]. 16 17 18 19 20 Dynesius, M. & Nilsson, C. (1994). Fragmentation and flow regulation of river systems in the northern third of the world. Science 266, 753-762. Ellenberg, H. (1988). Vegetation ecology of Central Europe. 4th edition. Cambridge Univ. Press., Cambridge, UK. European Environment Agency (2003). Europe’s environment: the third assessment. 21 Environmental Assessment Report No 10, Luxembourg: Office for Official 22 Publications of the European Communities. 6 1 Fischer, S., Poschlod, P. & Beinlich, B. (1996). Experimental studies on the dispersal of 2 plants and animals by sheep in calcareous grasslands. J. Appl. Ecol. 33, 1206- 3 1222. 4 Gijsbers, W.M. (1999). Kapitale ossen. De internationale handel in slachtvee in 5 Noordwest-Europa (1300-1750). PhD thesis University of Amsterdam. 6 [International trade in oxen in Northwest-Europe (1300-1750); In Dutch] 7 8 Gottschalk, M.K.E. 1977. Storm surges and river floods in the Netherlands. Social Geographical Studies 14, Assen. 9 Groot Bruinderink, G., van der Sluis, T., Lammertsma, D., Opdam, P. & Pouwels, R. 10 (2003). Designing a coherent ecological network for large mammals in 11 Northwestern Europe. Cons. Biol. 17, 549-557. 12 Hillegers, H.P.M. (1993). Heerdgang in Zuidelijk Limburg : een vorm van extensieve 13 beweiding in verleden, heden en toekomst. PhD thesis University of Maastricht. 14 [Livestock drifts in Southern Limburg, the Netherlands; In Dutch] 15 Hodgson, J.G. & Grime, J.P. (1990). The role of dispersal mechanisms, regenerative 16 strategies and seed banks in the vegetation dynamics of the British landscape. In: 17 Species Dispersal in Agricultural Habitats (eds. Bunce, R. G. H. & Howard, D. 18 C.) 65-82 (Belhaven Press, London. 19 20 21 22 Jager, S.W. (1985). A prehistoric route and ancient cart-tracks in the Gemeente of Anloo (Province of Drenthe). Palaeohistoria 27, 185-242. Jansson, R., Nilsson, C. & Renöfält, B. (2000). Fragmentation of riparian floras in rivers with multiple dams. Ecology 81, 899-903. 7 1 2 3 4 5 6 7 Janzen, D.H. & Martin, P.S. (1982). Neotropical anachronisms: The fruits the Gomphotheres ate. Science 215, 19-27. LWVT / SOVON. (2002). Bird migration in the Netherlands. Schuyt & Co, Haarlem. [in Dutch with English summary]. Lytle, D.A. & Poff, N.L. (2004). Adaptation to natural flow regimes. Trends Ecol. Evol. 19, 95-100. Kalweit, H., Ed. (1993). The Rhine under the influence of man. – River engineering 8 works, shipping, water management. Internationale Kommission für die 9 Hydrologie des Rheingebietes. Berichte Nr. I-11. [In German with English 10 11 12 13 summary]. KNMI (2005). Climate atlas of the Netherlands. Uitgeverij Elmar, Rijswijk. [in Dutch with English summary] Konold, W. (1997). Wässerwiesen, Wölbäcker, Hackäcker: Geschichte und Vegetation 14 alter Kulturlandschaftselemente in Südwestdeutschland. Verh. Ges. Ökologie 27, 15 53-61. [Flood meadows in SW Germany; in German]. 16 17 Manzano, P. & Malo, J.E. (2006). Extreme long-distance seed dispersal via sheep. Front. Ecol. Environm. 4, 244-248. 18 Myers, J.A., Vellend, M., Gardescu, S. & Marks, P.L. (2004). Seed dispersal by white- 19 tailed deer: implications for long-distance dispersal, invasion, and migration of 20 plants in eastern North America. Oecologia 139, 35-44. 21 Nathan, R. (2006). Long-distance dispersal of plants. Science 313, 786-788. 22 Nilsson, C., Reidy, C.A., Dynesius, M., Revenga, C. (2005). Fragmentation and flow 23 regulation of the world's large river systems. Science 308, 405-408. 8 1 2 3 4 5 6 Olff, H. & Ritchie, M.E. (1998). Herbivore effects on grassland plant species richness. Trends Ecol. Evol. 13, 261-265. Pakeman, R.J. (2001). Plant migration rates and seed dispersal mechanisms. J. Biogeogr. 28, 795-800. Poschlod, P., Bakker, J.P. & Kahmen, S. (2005). Changing land use and its impact on biodiversity. Basic Appl. Ecol. 6, 93-98. 7 Poschlod, P. & Bonn, S. (1998). Changing dispersal processes in the central European 8 landscape since the last ice age: an explanation for the actual decrease of plant 9 species richness in different habitats? Acta Bot. Neerl. 47, 27-44. 10 11 12 13 14 15 16 Pott, R. (1995). The origin of grassland plant species and grassland communities in Central Europe. Fitosociologia 29, 7-32. Rackham, O., (1986). The history of the countryside; the full fascinating story of Britain’s landscape. Dent, London. Ruiz, M. & Ruiz, J.P. (1986). Ecological history of transhumance in Spain. Biol. Conserv. 37, 73-86. Slichter Van Bath, B. H. (1980). De agrarische geschiedenis van West-Europa. 5th 17 edition. Uitgeverij Het Spectrum, Utrecht [The Agrarian History Of Western 18 Europe, A.D. 500-1850; In Dutch, but English translation by Olive Ordish 19 available for first edition 1963] 20 Spek, T. (2004). Het Drentse esdorpenlandschap: een historisch-geografische studie. 21 PhD thesis, University of Wageningen. Matrijs, Utrecht. [In Dutch with English 22 summary] 9 1 2 Stuart, A.J. (1991). Mammalian extinctions in the Late Pleistocene of northern Eurasia and North America. Biol. Rev. 66, 453-562. 3 Stuart, A.J. (2005). The extinction of woolly mammoth (Mammuthus primigenius) and 4 straight-tusked elephant (Palaeoloxodon antiquus) in Europe. Quaternary 5 International 126-28, 171-177. 6 Sutherland, G.D., Harestad, A.S., Price, K. & Lertzman, K.P. (2000). Scaling of natal 7 dispersal distance in terrestrial birds and mammals. Conserv. Ecol. 4, 16: 8 http://www.consecol.org/vol4/iss1/art16 9 10 11 12 13 14 15 Tockner, K. & Stanford, J.A. (2002). Riverine flood plains: present state and future trends. Env. Conserv. 29, 308-330. Wallis de Vries, M.F. (1995). Large herbivores and the design of large-scale nature reserves in Western-Europe. Conserv. Biol. 9, 25-33. Whittaker, C.R. (1988). Pastoral economies in classical antiquity. Cambridge Univ. Press, Cambridge. Wiese, H. (1966). Der Rinderhandel im nördwesteuropäischen Küstengebiet vom 15. 16 Jahrhundert bis zum Beginn des 19. Jahrhundert. In: H. Wiese & J. Bölts (eds.): 17 Rinderhandel und Rinderhaltung im nördwesteuropäischen Küstengebiet vom 15 18 Jahrhundert bis zum Beginn des 19 Jahrhundert. Stuttgart. [Cattle trade in coastal 19 areas of Northwest-Europe; In German] 20 10