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EARLY SUCCESSIONAL CHANGES IN FOREST ECOSYSTEMS AFTER A GAP-FORMING
DISTURBANCE
The project was supported by INTAS,
leader Prof. D.O. Logofet
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Project dates
2002-2005
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Summary
The Project was implemented during 2002-2005 in cooperation with the Chair of
Geobotany, M.V. Lomonosov Moscow State University, and western colleagues.
'Early successional changes in forest ecosystems after a gap-forming
disturbance' have been described as successive dynamics of species which
dominate in the lower forest layers inside the gap, i.e., the perennial grass
or/and shrub ones. The 'floristic relay' of these species depends
drastically on the forest type and the kind and strength of the disturbance.
In clear-cut areas of spruce forest (e.g., in the Bavarian Forest National
Park, Germany, or the Central Forest Reserve, Russia) the first colonizers
are red raspberry (Rubus idaeus) supplanted usually by small-reed of
Calamagrostis spp, an 'ideal weed' which is even able to block the tree
regeneration in an open gap; in windthrow areas of beech or spruce forest in
Swiss Alps, the Rubus spp. colonize quickly all the patches of disturbed soil
cover, then compete for light with survived undergrowth of tree spp. like
Betula pendula or Fagus sylvatica.
To understand the mechanisms underlying the interrelated dynamics of the first
colonizers, we analysed in the field, laboratories and computer rooms
the population structures in terms of the chronological age (in years)
and ontogenetic stages (along the scales developed formerly in the Russian
geobotanic school) of individual plants. Several sample plots were carefully
selected to represent certain stages of successional dynamics in three case
studies of the Project, and the experiments were laid down where each
individual plant, either survived one or appeared anew, has been marked
and monitored (once a year, in August) during the three-year Project
period. Certain parts of plants were dug out to provide for reliable
age determination in the laboratory, and the population age-stage
structures were established from the corresponding data-bases of
observations. The life cycle graphs (LCGs) have thereafter been
constructed as directed graphs of annual aging and ontogenetic
transitions over a grid of (a moderate number of) feasible
age-stage states. It has been found that plants respond to the
changing environment with changes in their ontogenetic pathways,
and the LCGs enable monitoring those changes from the year-to-year
observation data. The LCGs serve also as the conceptual bases for
mathematical modelling of the double-structured population dynamics.
A new family of matrix models for the double-structured populations has been
developed: the class of traditional Lefkovitch projection matrices for
stage-structured populations has been expanded (to Logofet-type nonnegative
matrices), the inherent mathematics has been investigated, and the
appropriate calibration procedures have been proposed and applied for the
particular case studies. When calibrated vs. the data, the projection
matrices have produced an estimate of the maximal population growth rate
1 (the dominant eigenvalue), the population structure that guarantees
the fastest growth (the corresponding dominant column-eigenvector), and
the vector (the corresponding dominant row-eigenvector) of
age-stage-specific reproductive values, i.e., the relative contributions
by the age-stage groups into the transient dynamics when the initial
population structure is different from that of the fastest growth.
These model findings have given quantitative certainty to theoretical
views on the population dynamics of colonizing spp., while the
(linear) matrix models have been found adequate to the phase of
extensive population growth.
However, after 5-10 years of the extensive growth, the first colonizer
undergoes competitor pressures or/and density self-regulation. To
illustrate the course of after-disturbance succession in the model terms,
we have modified the matrix models for double-structured, single-species
population dynamics of Rubus idaeus and Calamagrostis villosa into the
non-linear models which incorporate the density-dependent effects
responsible for various outcomes of the interrelated dynamics (i.e.,
competitive coexistence, suppression, or exclusion) at different phases
of the succession after clear cutting of spruce forest. A similar
approach can be recommended to model the outcome of succession under
various scenarios of forest management, which involve the gap-forming
disturbances.
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Publications
Logofet D.O., Ulanova N.G., Klochkova I.N., Demidova A.N.
Structure and dynamics of a clonal plant population:
Classical model results in a non-classic formulation.
Ecological Modelling,
2006, v.196, p. 95-106
Novoselova, E.A., Ulanova N.G., Fischer A.
The population structure of Rubus idaeus in beech-spruce
forests clearcuts in the Bavarian Forest National Park (Germany).
Topical Problems of Ecology and Conservancy.
2004, vol., 5-6, p.15-17. (in Russian).
Ulanova N.G., Klochkova I.N., Wohlgemuth Th., Logofet D.O.,
Nobis M.
An approach to combine dynamics of age and life stage in
clonal raspberry (Rubus idaeus L.) after disturbance.
Folia Geobotanica.
2007.
Ulanova, N.G., Zhukovskaya, O.V., Kuksina, N.V., Demidova, A.N.
Structure and dynamics of silver birch (Betula pendula Roth.)
in Calamagrostis epigeios communities on spruce clear-cuts
in Kostroma region.
Bull. Moscow Society of Naturalists, Biol. Section.
2005, v. 110, 5, p. 27-35.
Zhukovskaya O.V., Ulanova N.G.
Influence of brushing frequency on birch population
structure after felling.
Ecoscience.
2006, v.13, 2, pp. 219-225.
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Project participants
Laboratory of Mathematical Ecology (IAP RAS):
Dept. of Geobotany, Biological Faculty,
Lomonosov Moscow State University
Section of Geobotany, Dept. of Ecology, Center of Life and Food
Science, TU Munchen, Freising, Germany
WSL, Swiss Federal Institute for Forest, Snow and Landscape Research
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