Plant diversity in a managed temperate deciduous forest: understorey response to two silvicultural systems

Summary 1 Disturbances and resource availability are key factors affecting plant diversity in managed forests. As disturbance regimes vary among silvicultural systems and may simultaneously affect different types of resources, effects on biodiversity can be unpredictable. 2 We compared the effects of two silvicultural systems on understorey plant diversity, including species composition, structural attributes and functional organization. One hundred and thirty-five phytosociological releves were sampled from 27 forest stands managed under either a traditional coppice-with-standards (CWS, n = 12) or a ‘close-to-nature’ selective cutting system (SC, n = 15), over similar edaphic conditions. Important environmental factors affecting vegetation were deduced using Ellenberg indicator values. Structural diversity was described using traditional indices of α and β diversity. Guilds were defined within the local pool of species using a set of 14 traits and their relationship with silviculture was assessed using correspondence analysis. 3 Post-logged CWS stands share some compositional and structural characteristics with selectively cut stands, including high species richness and a dominance of early successional species. However the species pool for all coppicing areas was higher than for selectively cut areas, suggesting that the high disturbance frequency occurring in the latter may progressively eliminate the most sensitive species. 4 Functional diversity strongly differs between the two systems. Although it is conserved through the silvicultural cycle in the coppice-with-standards system, some guilds were lacking in selectively cut stands. The most negatively impacted guilds were tree and shrub saplings, prostrated ruderals, shade-tolerant perennials and vernal geophytes. The latter two comprise ‘true forest species’ which may also be considered as ‘coppicing-maintained species’. To reach the same values of guild richness (i.e. number of guilds) or redundancy (i.e. proportion of the maximal species richness within each guild), larger areas were required in SC compared with CWS systems. 5 In the SC system, the high proportion of light reaching the forest floor induced a spectacular spread of blackberries Rubus fruticosus agg., which decreased species richness. It also caused shifts in guild composition: graminoids and ferns grew strongly to the detriment of true forest species. 6 Synthesis and applications. Our results suggest long-term negative effects of selective cutting on both structural and functional plant diversity, compared with coppice-with-standards. Cutting intervals are shorter than recovery times, so that early successional species-dominated communities are maintained. Vernal geophytes and shade-tolerant perennials seem to be limited by the frequency of disturbance rather than by the severity of disturbance. We conclude that, from a biodiversity point of view, this ‘close-to-nature’ system does not cope with the objective of sustainable forest management. The rotations currently in use do not match natural disturbances very closely and are applied to a managed system rather than a natural forest. Retaining remnants of old coppice woods and extending rotations to at least 50 years are recommended where biodiversity conservation is a goal of forest management.

[1]  J A Drake,et al.  Divergent perspectives on community convergence. , 1997, Trends in ecology & evolution.

[2]  S. Lavorel,et al.  Predicting changes in community composition and ecosystem functioning from plant traits: revisiting the Holy Grail , 2002 .

[3]  G. W. Wendel,et al.  Plant Succession and Hydrologic Recovery on a Deforested and Herbicided Watershed , 1983 .

[4]  G. Decocq,et al.  Are there herbaceous dryads in temperate deciduous forests? , 2003 .

[5]  J. Barkman New systems of plant growth forms and phenological plant types. , 1988 .

[6]  R. H. Whittaker,et al.  Dominance and Diversity in Land Plant Communities , 1965, Science.

[7]  E. Maarel,et al.  Some remarks on disturbance and its relations to diversity and stability , 1993 .

[8]  M. Begon,et al.  Ecology: Individuals, Populations and Communities, 3rd edn. , 1997 .

[9]  Wolfgang Cramer,et al.  Plant functional types and disturbance dynamics – Introduction , 1999 .

[10]  M. Huston A General Hypothesis of Species Diversity , 1979, The American Naturalist.

[11]  Roderick Hunt,et al.  Comparative Plant Ecology: A Functional Approach to Common British Species , 1989 .

[12]  D. Tilman,et al.  Plant Dominance Along an Experimental Nutrient Gradient , 1984 .

[13]  Jean Thioulouse,et al.  ADE-4: a multivariate analysis and graphical display software , 1997, Stat. Comput..

[14]  J. Connell Diversity in tropical rain forests and coral reefs. , 1978, Science.

[15]  James H. Brown Two Decades of Homage to Santa Rosalia: Toward a General Theory of Diversity , 1981 .

[16]  J. Schultz,et al.  Understory Response to 50 Years of Management of a Northern Hardwood Forest in Upper Michigan , 1984 .

[17]  P. Grubb Plant Populations and Vegetation in Relation to Habitat, Disturbance and Competition: Problems of Generalization , 1985 .

[18]  J. Ehrenreich,et al.  Herbage production is related to hard-wood crown cover. , 1960 .

[19]  S. Lavorel,et al.  Plant functional classifications: from general groups to specific groups based on response to disturbance. , 1997, Trends in ecology & evolution.

[20]  E. Naumburg,et al.  Relationships between Pinus ponderosa forest structure, light characteristics, and understory graminoid species presence and abundance , 1999 .

[21]  G. Buckley,et al.  Ecology and management of coppice woodlands. , 1993 .

[22]  L. Firbank,et al.  An ecological comparison between ancient and other forest plant species of Europe, and the implications for forest conservation , 1999 .

[23]  G. Decocq Patterns of plant species and community diversity at different organization levels in a forested riparian landscape , 2002 .

[24]  D. Duffy,et al.  Do Appalachian herbaceous understories ever recover from clearcutting , 1992 .

[25]  G. Decocq,et al.  Soil seed bank composition and diversity in a managed temperate deciduous forest , 2004, Biodiversity & Conservation.

[26]  J. Denslow Patterns of plant species diversity during succession under different disturbance regimes , 1980, Oecologia.

[27]  M. Willig,et al.  The Relationship Between Productivity and Species Richness , 1999 .

[28]  K. Suding The effects of gap creation on competitive interactions: separating changes in overall intensity from relative rankings , 2001 .

[29]  D. Tilman,et al.  PLANT COMPETITION AND RESOURCE AVAILABILITY IN RESPONSE TO DISTURBANCE AND FERTILIZATION , 1993 .

[30]  A. Gillis The new forestry , 1990 .

[31]  Mats Hannerz,et al.  Effects on the flora in Norway spruce forests following clearcutting and shelterwood cutting , 1997 .

[32]  Charles B. Halpern,et al.  Plant Species Diversity in Natural and Managed Forests of the Pacific Northwest , 1995 .

[33]  J. Lawesson,et al.  Vegetation in Danish beech forests: the importance of soil, microclimate and management factors, evaluated by variation partitioning , 2004, Plant Ecology.

[34]  D. F. Grigal,et al.  INFLUENCE OF LOGGING, FIRE, AND FOREST TYPE ON BIODIVERSITY AND PRODUCTIVITY IN SOUTHERN BOREAL FORESTS , 2001 .

[35]  Professor Dr. Roelof A. A. Oldeman,et al.  Forests: Elements of Silvology , 1990, Springer Berlin Heidelberg.

[36]  M. Hunter,et al.  Natural disturbance as a guide to silviculture , 2002 .

[37]  Alain Franc,et al.  Biodiversity, disturbances, ecosystem function and management of European forests , 2000 .

[38]  Charles B. Halpern,et al.  PLANT DIVERSITY IN MANAGED FORESTS: UNDERSTORY RESPONSES TO THINNING AND FERTILIZATION , 1999 .

[39]  S. Bratton,et al.  Possible Ecological Mechanisms for Loss of Vernal‐Herb Diversity in Logged Eastern Deciduous Forests , 1995 .

[40]  B. Lacaze Annick Schnitzler-Lenoble, Ecologie des forêts naturelles d'Europe , 2002 .

[41]  P. Giller,et al.  Community structure and the niche , 1984 .

[42]  S. Lavorel,et al.  Plant life-history attributes: their relationship to disturbance response in herbaceous vegetation. , 1995 .

[43]  A. Schnitzler-Lenoble Ecologie des forêts naturelles d'Europe , 2002 .

[44]  B. Freedman,et al.  Forestry practices and biodiversity, with particular reference to the Maritime Provinces of eastern Canada , 1994 .

[45]  G. Decocq The ‘masking effect’ of silviculture on substrate-induced plant diversity in oak-hornbeam forests from northern France , 2000, Biodiversity & Conservation.

[46]  James B. Grace,et al.  The factors controlling species density in herbaceous plant communities: an assessment , 1999 .

[47]  M. Roberts,et al.  Early response of the herbaceous layer to harvesting in a mixed coniferous–deciduous forest in New Brunswick, Canada , 2002 .

[48]  E. D. Ford,et al.  The Biomass and Production of Ground Vegetation and Its Relation to Tree Cover Through a Deciduous Woodland Cycle , 1977 .

[49]  A. Kinzig,et al.  Original Articles: Plant Attribute Diversity, Resilience, and Ecosystem Function: The Nature and Significance of Dominant and Minor Species , 1999, Ecosystems.

[50]  N. Poff,et al.  Functional redundancy in heterogeneous environments: implications for conservation , 2001 .

[51]  Chapter 17 – Does intermediate disturbance increase species richness within deciduous forest understory? , 1991 .

[52]  M. L. Wolfe,et al.  Response of understory vegetation to variable tree mortality following a mountain pine beetle epidemic in lodgepole pine stands in northern Utah , 2004, Vegetatio.

[53]  Malcolm L. Hunter,et al.  Maintaining Biodiversity in Forest Ecosystems , 1999 .

[54]  R. Noss Indicators for Monitoring Biodiversity: A Hierarchical Approach , 1990 .

[55]  J. Anderson,et al.  Functional Ecology of Woodlands and Forests. , 1992 .

[56]  C. Braak,et al.  On the statistical analysis of vegetation change: A wetland affected by water extraction and soil acidification , 1994 .

[57]  C. Margules,et al.  Indicators of Biodiversity for Ecologically Sustainable Forest Management , 2000 .

[58]  D. Janzen Herbivores and the Number of Tree Species in Tropical Forests , 1970, The American Naturalist.

[59]  L. Taylor,et al.  The Ecology of Natural Disturbance and Patch Dynamics , 1986 .

[60]  R. Wein,et al.  Changes in understory vegetation with increasing stand age in New Brunswick forests: species composition, cover, biomass, and nutrients , 1977 .

[61]  T. Volpers Changes in microclimate and vegetation after thinning in a montane virgin forest , 1989 .

[62]  P. Grubb THE MAINTENANCE OF SPECIES‐RICHNESS IN PLANT COMMUNITIES: THE IMPORTANCE OF THE REGENERATION NICHE , 1977 .

[63]  David Tilman,et al.  Secondary Succession and the Pattern of Plant Dominance Along Experimental Nitrogen Gradients , 1987 .

[64]  M. Wassen,et al.  Calibrating Ellenberg indicator values for moisture, acidity, nutrient availability and salinity in the Netherlands , 1998, Plant Ecology.

[65]  W. Beese,et al.  Effect of alternative silvicultural systems on vegetation and bird communities in coastal montane forests of British Columbia, Canada , 1999 .

[66]  E. Bonnot,et al.  Nouvelle Flore de la Belgique, du Grand-Duche de Luxembourg, du Nord de la France et des Regions voisines , 1975 .

[67]  J. B. Faliński Vegetation Dynamics in Temperate Lowland Primeval Forests , 1986, Geobotany.

[68]  P. Alaback,et al.  Long-term response of understory vegetation to stand density in Picea-Tsuga forests , 1988 .

[69]  D. M. Newbery,et al.  M. A. Huston, Biological Diversity: the coexistence of species on changing landscapes . Cambridge University Press. ISBN 0-521-36930-4 (pbk). 681 + xix. pages. £24.95. , 1995, Journal of Tropical Ecology.

[70]  Steven L. Miller,et al.  Root Gap Dynamics in Lodgepole Pine Forest: Nitrogen Transformations in Gaps of Different Size , 1994 .

[71]  M. Willig,et al.  Randomness, Area, and Species Richness , 1982 .

[72]  D. Briske,et al.  Vegetation dynamics on rangelands: a critique of the current paradigms , 2003 .

[73]  W. Moir,et al.  Understory response to management treatments in northern Arizona ponderosa pine forests , 2001 .

[74]  J. P. Grime,et al.  Plant Strategies and Vegetation Processes. , 1980 .

[75]  E. van der Maarel,et al.  Transformation of cover-abundance values in phytosociology and its effects on community similarity , 1979, Vegetatio.

[76]  J. Bauhus C AND N MINERALIZATION IN AN ACID FOREST SOIL ALONG A GAP-STAND GRADIENT , 1996 .

[77]  B. Graae,et al.  The impact of forest continuity and management on forest floor vegetation evaluated by species traits. , 2000 .

[78]  David Tilman,et al.  The maintenance of species richness in plant communities , 1993 .