Seed limitation, not soil legacy effects, prevents native understorey from establishing in oak woodlands in Scotland after removal of Rhododendron ponticum

Following removal of the invasive species Rhododendron ponticum, the native understorey plant community typically fails to reestablish itself. Potential explanations for this failure include (1) lack of an appropriate native seed source; (2) inability of seed to penetrate a dense bryophyte layer; and (3) persistence of chemical “legacy effects” in the soil. We established an experiment to test these competing hypotheses in an Atlantic oak woodland where R. ponticum had been removed. The following experimental treatments were applied singly and in combination: (1) addition of a native seed mix to test for seed limitation; (2) removal of the established ground vegetation at the start of the experiment (which principally consisted of bryophytes) to test for the impact of a barrier layer; (3) addition of activated carbon to test for chemical legacy effects in the soil; and (4) fertilization as an additional measure to promote the establishment of native vascular plants. Application of the native seed mix was revealed to be an effective way to increase the cover of native vascular plants and was particularly effective when applied after the removal of the bryophyte layer. The application of activated carbon and/or fertilizer, however, had no effect on the cover of native vegetation. We conclude that reports of R. ponticum exerting chemical legacy effects long after its removal may have been overstated and that seed limitation and inability to successfully establish in a dense bryophyte layer provided the strongest barriers to natural recolonization by the native plant community following R. ponticum removal.

[1]  D. Burslem,et al.  Invasion by Rhododendron ponticum depletes the native seed bank with long-term impacts after its removal , 2018, Biological Invasions.

[2]  Forrest S. Smith,et al.  Comparing Three Common Seeding Techniques for Pipeline Vegetation Restoration: A Case Study in South Texas , 2015 .

[3]  V. Pillar,et al.  Are removal experiments effective tools for assessing plant community resistance and recovery from invasion , 2015 .

[4]  P. Pyšek,et al.  Impact of invasions by alien plants on soil seed bank communities: Emerging patterns , 2014 .

[5]  D. Richardson,et al.  Biological Invasions, resilience and restoration , 2012 .

[6]  C. D’Antonio,et al.  Gone but Not Forgotten? Invasive Plants' Legacies on Community and Ecosystem Properties , 2012, Invasive Plant Science and Management.

[7]  J. Ehrenfeld Ecosystem Consequences of Biological Invasions , 2010 .

[8]  K. Mganga,et al.  The challenges of rehabilitating denuded patches of a semi-arid environment in Kenya , 2010 .

[9]  B. Clinton,et al.  Variation in Soil and Forest Floor Characteristics Along Gradients of Ericaceous, Evergreen Shrub Cover in the Southern Appalachians , 2009 .

[10]  L. Morin,et al.  Does invasive plant management aid the restoration of natural ecosystems , 2009 .

[11]  K. Kiehl,et al.  Effects of a dense moss layer on germination and establishment of vascular plants in newly created calcareous grasslands , 2008 .

[12]  W. Bowman,et al.  Phenolic-rich leaf carbon fractions differentially influence microbial respiration and plant growth , 2008, Oecologia.

[13]  Dov F Sax,et al.  Species invasions and extinction: The future of native biodiversity on islands , 2008, Proceedings of the National Academy of Sciences.

[14]  Glenna M. Malcolm,et al.  Soil Nitrogen Conditions Approach Preinvasion Levels following Restoration of Nitrogen‐Fixing Black Locust (Robinia pseudoacacia) Stands in a Pine–Oak Ecosystem , 2008 .

[15]  R. Marrs,et al.  A restoration experiment on moorland infested by Pteridium aquilinum: Plant species responses , 2007 .

[16]  R. Hendrick,et al.  Rhododendron thickets alter N cycling and soil extracellular enzyme activities in southern Appalachian hardwood forests , 2007 .

[17]  C. Perrings,et al.  Controlling Rhododendron ponticum in the British Isles: an economic analysis. , 2004, Journal of environmental management.

[18]  G. Peterken Ecological effects of introduced tree species in Britain , 2001 .

[19]  R. Abbott,et al.  Origin and evolution of invasive naturalized material of Rhododendron ponticum L. in the British Isles , 2000, Molecular ecology.

[20]  R. Marrs,et al.  Vegetation re-establishment on land previously subject to control of Pteridium aquilinum by herbicide , 2000 .

[21]  O. Zackrisson,et al.  Characterisation of the differential interference effects of two boreal dwarf shrub species , 2000, Oecologia.

[22]  M. Zamfir Effects of bryophytes and lichens on seedling emergence of alvar plants: evidence from greenhouse experiments , 2000 .

[23]  B. Clinton,et al.  Inhibition of seedling survival under Rhododendron maximum (Ericaceae): could allelopathy be a cause? , 1999, American journal of botany.

[24]  M. Kent,et al.  Seed bank composition and variability in five woodlands in south-west England , 1994 .

[25]  M. Usher,et al.  Impact of carpets of the invasive moss Campylopus introflexus on Calluna vulgaris regeneration , 1993 .

[26]  D. Barker,et al.  Natural reseeding of five grass species in summer dry hill country , 1991 .

[27]  J. A. August,et al.  Use of nitrogen fertiliser in restoration of pasture productivity and soil fertility after topsoil mining , 1988 .

[28]  J. R. Cross Biological flora of the British Isles. Rhododendron ponticum L. , 1975 .

[29]  S. Meyer,et al.  Cheatgrass die-offs as an opportunity for restoration in the Great Basin, USA: Will local or commercial native plants succeed where exotic invaders fail? , 2016 .

[30]  R Core Team,et al.  R: A language and environment for statistical computing. , 2014 .

[31]  Shakhnoza S. Azimova,et al.  Rhododendron ponticum L. , 2012 .

[32]  C.J.F. ter Braak,et al.  Canoco Reference Manual and User’s Guide: Software for Ordination (version 5.0) , 2012 .

[33]  N. Dise,et al.  Consequences of lime and fertiliser application for moorland restoration and carbon balance , 2007 .

[34]  A. Scalbert,et al.  Polyphenols of Quercus robur: Adult tree and in vitro grown calli and shoots , 1988 .

[35]  D. Read,et al.  Aspects of the ecology of Rhododendron ponticum with reference to its competitive and invasive properties. , 1988 .

[36]  A. Scalbert,et al.  Polyphenols and chemical defence of the leaves of Quercus robur , 1987 .

[37]  I. Rotherham The ecology of Rhododendron ponticum L. with special reference to its competitive and invasive capabilities , 1983 .