Stand dieback and collapse in a temperate forest and its impact on forest structure and biodiversity

Abstract Concern is increasing about large-scale dieback that is occurring in many forest ecosystems. However, understanding of the processes of dieback and its potential impacts is limited, partly owing to the lack of long-term monitoring data for forest stands in which dieback has occurred. Here we present monitoring data collected over 50 years along two transects in a temperate forest ecosystem, in which the canopy dominant beech (Fagus sylvatica L.) has demonstrated significant dieback. Our results show that basal area in the forest has declined by 33%, and juvenile tree densities have also been reduced by approximately 70%. Growing season temperatures have steadily increased and there have been a number of droughts causing climatic water deficits in recent decades, particularly in 1995. We hypothesise that these droughts may have interacted with novel pathogenic fungi to cause mortality of large trees. Curvilinear responses to BA loss were observed in tree community change, ground flora species richness, and percentage cover of grass, providing evidence of thresholds associated with stand dieback. Evidence also suggested that BA failed to recover once it declined. Critical values of basal area for a change in ground flora species richness and grass cover were around 40% decline from initial values. While these changes are dramatic, they cannot be considered a regime shift as the pressures that may have contributed to the ecosystem transition, drought, pathogenic fungi and overgrazing, are on-going. While managers might consider accepting forest dieback as part of an adaptive response of the system to novel environmental conditions, this would likely be associated with significant change in biodiversity and ecosystem service provision.

[1]  T. Jung,et al.  Beech decline in Central Europe driven by the interaction between Phytophthora infections and climatic extremes , 2009 .

[2]  C. Leuschner,et al.  Belowground drought response of European beech: fine root biomass and carbon partitioning in 14 mature stands across a precipitation gradient , 2008 .

[3]  A. Newton,et al.  Dynamics and Conservation Management of a Wooded Landscape under High Herbivore Pressure , 2013 .

[4]  Werner A. Kurz,et al.  Risk of natural disturbances makes future contribution of Canada's forests to the global carbon cycle highly uncertain , 2008, Proceedings of the National Academy of Sciences.

[5]  S. Carpenter,et al.  Anticipating Critical Transitions , 2012, Science.

[6]  Paul C. Johnson Extension of Nakagawa & Schielzeth's R2GLMM to random slopes models , 2014, Methods in ecology and evolution.

[7]  R. Finkeldey,et al.  Are beech (Fagus sylvatica) roots territorial , 2010 .

[8]  G. Peterken,et al.  Effects of drought on beech in Lady Park Wood, an unmanaged mixed deciduous woodland , 1996 .

[9]  A. Sala,et al.  Physiological mechanisms of drought-induced tree mortality are far from being resolved. , 2010, The New phytologist.

[10]  A. Taylor,et al.  Widespread Increase of Tree Mortality Rates in the Western United States , 2009, Science.

[11]  Adrian C. Newton,et al.  Restoration of forest resilience: An achievable goal? , 2015, New Forests.

[12]  David R. Anderson,et al.  AIC model selection and multimodel inference in behavioral ecology: some background, observations, and comparisons , 2011, Behavioral Ecology and Sociobiology.

[13]  Climate Warming-Related Growth Decline Affects Fagus sylvatica, But Not Other Broad-Leaved Tree Species in Central European Mixed Forests , 2015, Ecosystems.

[14]  Jerry F. Franklin,et al.  Climatic water deficit, tree species ranges, and climate change in Yosemite National Park , 2010 .

[15]  I. Boyd,et al.  The Consequence of Tree Pests and Diseases for Ecosystem Services , 2013, Science.

[16]  A. Nardini,et al.  Global convergence in the vulnerability of forests to drought , 2012, Nature.

[17]  M. Manthey,et al.  Drought matters – Declining precipitation influences growth of Fagus sylvatica L. and Quercus robur L. in north-eastern Germany , 2011 .

[18]  S. Carpenter,et al.  Catastrophic shifts in ecosystems , 2001, Nature.

[19]  Marten Scheffer,et al.  Resilience thinking: integrating resilience, adaptability and transformability , 2010 .

[20]  P. Petraitis,et al.  Multiple Stable States in Natural Ecosystems , 2013 .

[21]  Adrian C. Newton,et al.  Social-ecological Resilience and Biodiversity Conservation in a 900-year- old Protected Area , 2011 .

[22]  Giovanni Coco,et al.  Forecasting the limits of resilience: integrating empirical research with theory , 2009, Proceedings of the Royal Society B: Biological Sciences.

[23]  Shinichi Nakagawa,et al.  A general and simple method for obtaining R2 from generalized linear mixed‐effects models , 2013 .

[24]  O. Holdenrieder,et al.  European ash (Fraxinus excelsior) dieback - a conservation biology challenge. , 2013 .

[25]  Hadley Wickham,et al.  ggplot2 - Elegant Graphics for Data Analysis (2nd Edition) , 2017 .

[26]  David D. Briske,et al.  A Unified Framework for Assessment and Application of Ecological Thresholds , 2006 .

[27]  Wolfgang Lucht,et al.  Forest resilience and tipping points at different spatio‐temporal scales: approaches and challenges , 2015 .

[28]  Christof Bigler,et al.  Drought induces lagged tree mortality in a subalpine forest in the Rocky Mountains , 2007 .

[29]  J. Hicke,et al.  Cross-scale Drivers of Natural Disturbances Prone to Anthropogenic Amplification: The Dynamics of Bark Beetle Eruptions , 2008 .

[30]  S. Jørgensen Model Selection and Multimodel Inference: A Practical Information—Theoretic Approach, Second Edition, Kenneth P. Brunham, David R. Anderson, Springer-Verlag, Heidelberg, 2002, 490 pages, hardbound, 31 illustrations , 2004 .

[31]  Kenneth A. Dawson,et al.  Role of cell cycle on the cellular uptake and dilution of nanoparticles in a cell population. , 2011, Nature nanotechnology.

[32]  A. Newton Biodiversity in the New Forest , 2010 .

[33]  Lael Parrott,et al.  From Management to Stewardship: Viewing Forests As Complex Adaptive Systems in an Uncertain World , 2015 .

[34]  S. Carpenter,et al.  Catastrophic regime shifts in ecosystems: linking theory to observation , 2003 .

[35]  C. Peng,et al.  A drought-induced pervasive increase in tree mortality across Canada's boreal forests , 2011 .

[36]  Frank Fleischmann,et al.  Involvement of Phytophthora species in the decline of European beech in Europe and the USA , 2006 .

[37]  Sebastiaan Luyssaert,et al.  Carbon sequestration: Managing forests in uncertain times , 2014, Nature.

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

[39]  B. Pedersen,et al.  THE ROLE OF STRESS IN THE MORTALITY OF MIDWESTERN OAKS AS INDICATED BY GROWTH PRIOR TO DEATH , 1998 .

[40]  Francis K C Hui,et al.  The arcsine is asinine: the analysis of proportions in ecology. , 2011, Ecology.

[41]  B. Pedersen THE MORTALITY OF MIDWESTERN OVERSTORY OAKS AS A BIOINDICATOR OF ENVIRONMENTAL STRESS , 1999 .

[42]  M. Lesperance,et al.  PIECEWISE REGRESSION: A TOOL FOR IDENTIFYING ECOLOGICAL THRESHOLDS , 2003 .

[43]  N. McDowell,et al.  The interdependence of mechanisms underlying climate-driven vegetation mortality. , 2011, Trends in ecology & evolution.

[44]  J. Peñuelas,et al.  Rapid climate change‐related growth decline at the southern range edge of Fagus sylvatica , 2006 .

[45]  Scott J. Goetz,et al.  Terrestrial and Inland Water Systems , 2014 .

[46]  J. Healey,et al.  Evaluation of Forest Recovery over Time and Space Using Permanent Plots Monitored over 30 Years in a Jamaican Montane Rain Forest , 2012, PloS one.

[47]  G. Peterken,et al.  Long‐term change and implications for the management of wood‐pastures: experience over 40 years from Denny Wood, New Forest , 2003 .

[48]  Adrian C. Newton,et al.  Identifying cost-effective indicators to assess the conservation status of forested habitats in Natura 2000 sites , 2008 .

[49]  M. Rietkerk,et al.  Self-Organized Patchiness and Catastrophic Shifts in Ecosystems , 2004, Science.

[50]  N. McDowell,et al.  A global overview of drought and heat-induced tree mortality reveals emerging climate change risks for forests , 2010 .

[51]  C. S. Holling,et al.  Resilience, Adaptability and Transformability in Social–ecological Systems , 2004 .

[52]  Sabine Augustin,et al.  The Scientific Potential of Environmental Monitoring , 2010 .

[53]  Todd E. Ristau,et al.  Chronic over browsing and biodiversity collapse in a forest understory in Pennsylvania: Results from a 60 year-old deer exclusion plot , 2011 .

[54]  David R. Anderson,et al.  Model selection and multimodel inference : a practical information-theoretic approach , 2003 .

[55]  G. F. Peterken,et al.  Long-term change in growth, mortality and regeneration of trees in Denny Wood, an old-growth wood-pasture in the New Forest (UK) , 1999 .

[56]  P. Thomas,et al.  Biological Flora of the British Isles: Fagus sylvatica , 2012 .

[57]  A. Huggett The concept and utility of ecological thresholds in biodiversity conservation , 2005 .

[58]  Monica G. Turner,et al.  Ecological Thresholds: The Key to Successful Environmental Management or an Important Concept with No Practical Application? , 2006, Ecosystems.

[59]  D. Bates,et al.  Linear Mixed-Effects Models using 'Eigen' and S4 , 2015 .