Succession theory and wetland restoration ecology
Wu Xinyuan
(Texas
A&M University, College Station, TX 77843-2126, USA)
Abstract:
Succession theory, as the basis for prediction of wetland
restoration, plays an important role in wetland restoration
ecology. Indeed,
ecological restoration has been considered as acceleration of
succession. Wetlands
have been fertile ground for studies of ecological succession over
the last century. Most
of these empirical studies on community dynamics in wetlands yield
evidences that are considered in support of the Gleasonian and not
the Clementsian succession theory. The contrasting views of the Clementsian and Gleasonian also
have been involved often in the discussion of the theories of
ecological restoration. This
paper attempts to examine the linkage between historical
succession theories and current schools of thoughts in wetland
restoration ecology and discuss the need and directions to broaden
the theoretical framework for the development of wetland
restoration ecology.
There have been two major
schools of thoughts on wetland restoration ecology, self-design
vs. designer-design. The
self-design approach takes more an ecosystem approach and
emphasize that the nature of the environment (including the
disturbance regimes) and the ecosystem processes dictates the
composition and spatial pattern of plants in wetlands.
Self-designers question the necessity and long-term effectiveness
of the human intervention of designed planting in hydrologically
open wetland systems and believe plant species will be sort out
spatially when prorogues are available. They stress that the
development of ecosystem functions may take considerable time and
human intervention can help accelerate their development. The designer-design approach takes more a reductionist,
population-level approach and emphasizes the importance of
designed planting based on sound understanding of life history
attributes of the species involved.
Designer proponents believe, because the lack of sufficient
seed bank and limited dispersal, as well as the strong influence
of initial floristic composition in many cases, human intervention
is critical in determine the eventual outcome of restored
wetlands.
Some scholars have argued that
the self-design and designer theories are simply restatements of
the Clementsian and Gleasonian succession theories.
This characterization, however, is not quite accurate.
Although influenced by Odum’s ecosystem ecology, the
self-design theory does not include some of the basic elements of
Clementsian theory, such as a predictable, orderly process toward
a stable climatic climax. Its
strong emphasis on the role of the physical environment and
disturbance regime in sorting the composition and spatial
distribution of species actually appears more Gleasonian than
Clementsian. The
emphasis of the designer approach on population-level processes as
mechanisms of community dynamics is Gleasonian and is essential to
the understanding and development of wetland restoration.
Gleasonian emphasis on the role of chance events, however,
can be for or against the key element of designer theory, designed
planting; planting can prevent the development of unwanted
structure due to chance event such as preemption of undesirable
species, but the long-term effects of planting can be reduced or
overridden by disturbances that are stochastic in nature.
The different emphases of the
self-design and designer theories are influenced to a degree by
different goals of wetland restoration, self-design tends to focus
more on ecosystem function and less on biodiversity than designer.
The goal setting of self-design tends to focus on longer
temporal scale than that of designer as well.
Experiences with different wetland ecosystem of the
proponents of the two theories may have also played a role.
The designer thoughts are strongly linked to studies of
prairie potholes where seed bank is a key element of the wetland
dynamics and dispersal is often limited in the agriculture
landscape with low connectivity among wetlands.
The self-design thoughts are strongly linked to studies of
hydrologically open wetlands such as riparian systems where
availability and dispersal of prorogues are not limiting.
Neither self-design nor designer theory appears have expanded
sufficiently beyond the traditional succession theories and
explicitly incorporated some of the contemporary views of
vegetation dynamics. A
more comprehensive conceptual framework that integrates key
elements of both self-design and designer theories and explicitly
anchors on contemporary theories of vegetation dynamics is needed
for the development of wetland restoration ecology.
Several theoretical issues key to the development of
wetland restoration ecology include the following.
(1) Adopt the view that continuous change in vegetation is
the norm (similar to Cowles’ view of succession) and that
multiple steady states exist in ecosystem dynamics.
This may require fundamental modifications of our concepts
of reference conditions (e.g., multiple reference states and
contingency/redundancy) and restoration designs.
(2) Explicitly address spatial and temporal scales of
disturbance regimes and the development of different ecosystem
structures and functions. Development
of some ecosystem attributes, such as soil development and
nutrient cycle, are critical factors influencing community
development but can take considerable amount of time.
This may not only affect expected timeframe for restoration
but also the specificity of goals of restoration given the
stochastic nature of disturbances events and other long-term
(e.g., climatic) changes. (3)
Emphasize the landscape context for wetland restoration.
More than those in other areas of ecological restoration,
wetland restorationists appreciate the importance of landscape
context likely due to their awareness of the inherently critical
importance of hydrologic regimes and plant dispersal to wetland
restoration; a stronger emphasis, however, is the need on
quantitative assessment of landscape setting and dynamics and
their influence to the dynamics and long-term fate of restored
wetlands, as well as the expected distribution and function of the
wetlands in the watershed. (4)
Recognize the potential limitations of succession theories and
empirical knowledge for restoration applications because the
significant differences between restoration and natural
succession. Restoration
starts with an unnatural setting with often sudden changes in the
environmental conditions and artificially synchronized
introduction of vegetation, which likely invoke spatial and
temporal patterns of interactions and processes different from
those operate in natural succession; in addition, some of the
basic processes, such as hydrology/flooding regime in urbanized
watersheds, have been fundamentally altered by human disturbances,
which may result in novel settings that can induce new patterns
and mechanisms of ecosystem dynamics.
(5) Promote the incorporation of large-scale experiments
into restoration projects. This
may allow testing and developing new succession and restoration
theories, which is essential to the development of the science and
practice of restoration ecology.
Key words: wetland restoration ecology; succession theory; self-design
vs.designer-design
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作者简介:武昕原(Xinyuan
Ben Wu),男,1956年生,美国田纳西州大学生态学博士,现就任于德克萨斯A&M大学,副教授。研究方向为湿地生态与恢复、景观生态学、保护生态学、自然资源管理。主持研究美国科学基金的稀树草原碳氮储量测度、美国国家公园服务局的城市化景观的湿地恢复、德克萨斯A&M大学的基于GIS的城市景观生态学研究途径等项目,发表论文40多篇。Email: b-wu@tamu.edu
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