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Ancient and recent alien species in temperate forests: steady state and time lags

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Abstract

Temperate forests are relatively little affected by invasions globally. However, it remains unclear if these low invasion levels can be attributed to high invasion resistance, or if particularly long time lags in forests have lead to wide-spread invasion debt at the landscape and habitat scales, thus masking the invasibility of forests. Using linear and generalized linear mixed-effects models, we tested which factors determine the level of invasion of old (archaeophytes, pre-1500 alien species) and new (neophytes, post-1500 alien species) invaders across 30 study sites (5 ha size) and 16 transects in lowland forests in eastern Austria, Central Europe. We analysed the importance of different forest types, adjacent habitats, introduction pathways, species’ local residence time and proximity to human settlements as proxy for propagule pressure. We recorded 109 neophyte and 50 archaeophyte species, with significant differences between both groups with respect to region of origin, pathways, and life form. The number of neophyte species per study site varied by more than an order of magnitude and was dependent on both proximity to settlements and residence time in a non-linear manner, with proximity to settlements showing a particularly strong effect for recently introduced neophytes. In contrast, archaeophyte species numbers were only marginally affected by distance to the nearest settlement. Within habitats, transect plots close to the forest edge and adjacent to settlements showed significantly higher numbers and cover of neophyte species, whereas for archaeophytes and native species no such effect could be detected. Our results suggest that time lag phenomena stemming from dispersal limitation (e.g. aggravated by isolation of forest patches), heterogeneity in species’ local residence time as well as increases in local levels of propagule pressure (e.g. created by expanding settlements), may result in wide-spread invasion debt at the landscape and habitat scales, hence masking invasibility of forests. Spread of new invaders may ultimately, although possibly only over long time periods, result in increased frequency and impact of neophyte species in temperate forests.

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Acknowledgments

We are grateful to Christian Schröck and Oliver Stöhr for their assistance in determining critical specimens. We greatly appreciate comments from the associate editor, Marc Cadotte, and from two anonymous reviewers, which helped to improve the manuscript.

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Authors and Affiliations

  1. Department for Biodiversity and Nature Conservation, Environment Agency Austria, Spittelauer Lände 5, 1090, Vienna, Austria

    Franz Essl & Dietmar Moser

  2. Vienna Institute for Nature Conservation and Analyses, Giessergasse 6/7, 1090, Vienna, Austria

    Thomas Mang & Dietmar Moser

Authors
  1. Franz Essl
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  2. Thomas Mang
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  3. Dietmar Moser
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Corresponding author

Correspondence to Franz Essl.

Electronic supplementary material

Below is the link to the electronic supplementary material.

10530_2011_156_MOESM1_ESM.doc

Appendix S1: Forest types (in % of total area), identity of adjacent habitat types (1 = present, 0 = absent) and total number of adjacent habitat types, minimum distance to the next settlement, and total neophyte and archaeophyte species numbers in the 30 study sites. Abbreviations: native broadl. for. = native broadleaved forests dominated by oaks (Quercus cerris, Q. petraea, Q. robur); conifer for. = native conifer plantations (Pinus nigra, P. sylvestris); alien broadl. for. = forests dominated by alien broadleaved trees (Ailanthus altissima, Robinia pseudacacia). Supplementary material 1 (DOC 95 kb)

10530_2011_156_MOESM2_ESM.doc

Appendix S2: Neophyte flora of the 30 study sites. Given are taxon name, year of first record in Austria (year), pathway (hort = horticulture, for = forestry, unint = unintentional), origin (NA = North America, AS = Asia, cult = originated in horticulture - anecophytes, EU = Europe), life form (ll = long-lived herb, tr = tree, sh = shrub, sl = short-lived herb). Occurrence in the forest patches is recorded in five logarithmic abundance classes (VC = very common, i.e. >10% plant cover; CO = common, i.e. 1-10% plant cover; SC = scattered, i.e. 0.1-1% plant cover; RA = rare, i.e. few individuals with 0.1-0.01% plant cover; VR = very rare, i.e. very few individuals with very low plant cover, <0.01) and total number of colonized sites (no). Supplementary material 2 (DOC 610 kb)

10530_2011_156_MOESM3_ESM.doc

Appendix S3: Archaeophyte flora of the 30 study sites. Given are taxon name, pathway (hort = horticulture, agri = agriculture, unint = unintentional), origin (EU = Europe), life form (ll = long-lived herb, tr = tree, sh = shrub, sl = short-lived herb). Occurrence in the forest patches is recorded in five logarithmic abundance classes (VC = very common, i.e. >10% plant cover; CO = common, i.e. 1-10% plant cover; SC = scattered, i.e. 0.1-1% plant cover; RA = rare, i.e. few individuals with 0.1-0.01% plant cover; VR = very rare, i.e. very few individuals with very low plant cover, <0.01) and total number of colonized sites (no). Supplementary material 3 (DOC 283 kb)

10530_2011_156_MOESM4_ESM.doc

Appendix S4: Plant species data of the 16 transects. Each transect consists of five plots, which are labelled one to five with increasing distance from the forest edge. Plant species cover is given in %, cover is assessed separately for herb, shrub and tree layers. Further, floristic status (native, archaeophyte, neophyte), residence time in Austria, the dominant pathway, region of origin, and the life form are given. Abbreviations: TL = tree layer, SL = shrub layer, HL = herb layer. Supplementary material 4 (DOC 159 kb)

10530_2011_156_MOESM5_ESM.doc

Appendix S5: The linear mixed-effects models (LMMs) for alien (archaeophytes, neophytes) (A) and native (B) species total plant cover in the transect plots. Analyses were restricted to species present at each plot. Estimated standard deviations of random-effect intercepts and residual term: alien species: transect: 0.15, plot: < 0.01, species: 0.57, residual: 0.82; native species: transect: 0.15, plot: < 0.01, species: 0.55, residual: 0.82. Supplementary material 5 (DOC 43 kb)

10530_2011_156_MOESM6_ESM.doc

Appendix S6: The linear mixed-effects models (LMMs) for alien (archaeophytes, neophytes) (A) and native (B) species herb-layer cover in the transect plots. Analyses were restricted to species present at each plot. Standard deviations of random-effect intercepts and residual term: alien species: transect: 0.07, plot: < 0.01, species: 0.45, residual: 0.70; native species: transect: 0.09, plot: < 0.01, species: 0.46, residual: 0.64. Supplementary material 6 (DOC 42 kb)

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Essl, F., Mang, T. & Moser, D. Ancient and recent alien species in temperate forests: steady state and time lags. Biol Invasions 14, 1331–1342 (2012). https://doi.org/10.1007/s10530-011-0156-y

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