Connell–Slatyer model of ecological succession

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Ecological succession can be understood as a process of changing species composition within a community due to an ecological disturbance, and varies largely according to the initial disturbance prompting the succession. [1] Joseph Connell and Ralph Slatyer further developed the understanding of successional mechanisms in their 1977 paper and proposed that there were 3 main modes of successional development. These sequences could be understood in the context of the specific life-history theories of the individual species within an ecological community.

Contents

The 1977 study

Connell and Slatyer chose to focus on autogenic succession, which occurs on newly exposed landforms and is initiated by changes from within the community rather than a geophysical transformation. [2] They targeted plant and immobile aquatic organisms that demanded the greatest surface area within an environment and could modify the physical landscape. They defined community as "the set of organisms that occur together and that significantly affect each other's distribution and abundance".

The models

The key factor distinguishing the three models is how the process of succession affects the original, pioneer species (i.e. their relative success in later-successional stages). [3]

Facilitation model

Facilitation Model Facilitation.JPG
Facilitation Model

Based on the assumption that only particular species with qualities ideal for "early succession" can colonize the newly exposed landforms after an ecological disturbance.

  • These "colonizing" qualities include: highly effective methods of dispersal, the ability to remain dormant for long periods of time, and a rapid growth rate. However, the pioneer species are often subsequently less successful once an area has been heavily populated by surrounding species due to increased shade, litter or concentrated roots in the soil, etc. [3]
  • Thus, the presence of early successional species often changes the environment so that the habitat is less hospitable for the original species’ own ecological demands and facilitates the invasion of later-successional species. [3] (Note: See Ecological Facilitation.)

Tolerance model

Tolerance Model Tolerance.JPG
Tolerance Model

In this case, new pioneer species neither inhibit nor facilitate the growth and success of other species. The sequences of succession are thus entirely dependent on life-history characteristics such as the specific amount of energy a species allocates to growth. [3]

Inhibition model

Inhibition Model Inhibition.JPG
Inhibition Model

Earlier successional species actually inhibit growth of later successional species and reduce growth of colonizing species already present. [3]

  • Example: Pioneer species might modify the environment through rapid growth and make the area increasingly shady (essentially increasing competition for light). [3]
  • The environment is thus less hospitable to other potential colonizing species. [3]
  • The only possibility for new growth/colonization in this successional sequence arises when a disturbance leads to dominating species being destroyed, damaged, or removed. This frees up resources and allows for the invasion of other species that were not previously present. [3]

Examples of each model

1. Facilitation Model
Essentially, the facilitation model suggests that the presence of an initial species aids and increases the probability of the growth of a second species. [4] For example, in "Physiological Controls Over Seedling Growth in Primary Succession on an Alaskan Floodplain" by Lawrence R. Walker and F. Stuart Chapin, III, the presence of alder plants aids the growth of willow and poplar seedlings in an Alaskan floodplain. [5]
Alder roots contain nitrogen-fixing bacteria, which greatly increase the amount of inorganic nitrogen present in soils. [6] This increased availability of nitrogen aids the growth of both willow and poplar seedlings in areas without other competition. Eventually, however, willow and poplar grow more rapidly than alder, leading to a reduction in the abundance of the pioneer species, and eventually, spruce becomes a later-succession species, due to its increased ability (over alder) to grow in shaded areas. [7]
Another case of facilitation comes from the colonization of lakeshore sand dunes. [8] Adjacent pioneer plants colonize the otherwise moving sands and alter the environmental constraints of the sandy environment to better suit other plant species, which can then allow for soil binding to take place. [9] The giant saguaro cactus, in this respect, can only survive in the shade of other plants (or in some cases rocks) – pioneer species facilitate their existence by providing shade. [8] (The argument has also been made that this type of interaction is exemplary of the tolerance model; see below). [8]
2. Tolerance Model
The tolerance model is completely dependent upon life history characteristics. Each species has an equally likely chance to establish itself in the early stages of succession and their establishment results in no environmental changes or impacts on other species. [10] Eventually, early species, typically dominated by r-selected species, which prioritize fast rates of reproduction, are out-competed by K-selected species (species that become more dominant when there is competition for limited resources). [10]
For example, we can examine succession in the Loess Plateau in China. In the graph on page 995 of the paper "Plant Traits and Soil Chemical Variables During a Secondary Vegetation Succession in Abandoned Fields on the Loess Plateau" by Wang (2002), we can see the initial dominance of the Artemisia scoparia , the pioneer species. Over time, however, the Bothriochloa ischaemum becomes the dominant species and the abundance of A. scoparia greatly declines. This is due to the rapid rate of reproduction of the A. scoparia, resulting in the species' early abundance, and the dominant competition from the K-selected B. ischaemum, resulting in that species' later abundance. [11]
A characteristic that is often associated with the tolerance model and well documented in forest succession is survival in conditions of shade. [12] As an uninhabited area becomes populated by different plant species, shade increases – which makes less light available for the next generation. Species that are better adapted to shady conditions will then become dominant. All of the early species inhabiting the terrain have modified the environment in a way that favors a specific k-selected characteristic. [2]
3. Inhibition Model
In this model, one species inhibits the presence of another, either through direct means, such as predation (by eating the other species or attacking them), or indirect means, such as competition for resources. [4]
Sometimes in inhibition models, the time of establishment of a species determines which species becomes dominant. This phenomenon is referred to as the priority effect and suggests that the species that became established earlier are more likely to become the dominant species. One example of the inhibition model, and the priority effect, occurs in South Australia. In areas where bryozoans are established first, tunicates and sponges cannot grow. [13]
The inhibition model has also been observed at work in forest ecosystems; in these systems the early arrivers hold a monopoly on the land, keeping other species out. Closed shrub canopies have been known to prevent tree growth and access to land for periods of up to 45 years – in an experimental study on inhibition it was found that areas occupied by large areas of Lantana sprawling shrubs excluded and inhibited the growth of tree species. [14]

Related Research Articles

Ecological selection refers to natural selection without sexual selection, i.e. strictly ecological processes that operate on a species' inherited traits without reference to mating or secondary sex characteristics. The variant names describe varying circumstances where sexual selection is wholly suppressed as a mating factor.

<span class="mw-page-title-main">Pioneer species</span> First species to colonize or inhabite damaged ecosystems

Pioneer species are hardy species that are the first to colonize barren environments or previously biodiverse steady-state ecosystems that have been disrupted, such as by wildfire.

<span class="mw-page-title-main">Climax community</span> Ecological community of organisms

In scientific ecology, climax community or climatic climax community is a historic term for a community of plants, animals, and fungi which, through the process of ecological succession in the development of vegetation in an area over time, have reached a steady state. This equilibrium was thought to occur because the climax community is composed of species best adapted to average conditions in that area. The term is sometimes also applied in soil development. Nevertheless, it has been found that a "steady state" is more apparent than real, particularly across long timescales. Notwithstanding, it remains a useful concept.

<span class="mw-page-title-main">Ecological succession</span> Process of change in the species structure of an ecological community over time

Ecological succession is the process of change in the species structure of an ecological community over time. The time scale can be decades or more or less.

<span class="mw-page-title-main">Primary succession</span> Gradual growth and change of an ecosystem on new substrate

Primary succession is the beginning step of ecological succession after an extreme disturbance, which usually occurs in an environment devoid of vegetation and other organisms. These environments are typically lacking in soil, as disturbances like lava flow or retreating glaciers shred the environment of nutrients.

<span class="mw-page-title-main">Forest dynamics</span>

Forest dynamics describes the underlying physical and biological forces that shape and change a forest ecosystem. The continuous state of change in forests can be summarized with two basic elements: disturbance and succession.

<span class="mw-page-title-main">Intermediate disturbance hypothesis</span> Model proposing regional biodiversity is increased by a moderate level of ecological disturbance

The intermediate disturbance hypothesis (IDH) suggests that local species diversity is maximized when ecological disturbance is neither too rare nor too frequent. At low levels of disturbance, more competitive organisms will push subordinate species to extinction and dominate the ecosystem. At high levels of disturbance, due to frequent forest fires or human impacts like deforestation, all species are at risk of going extinct. According to IDH theory, at intermediate levels of disturbance, diversity is thus maximized because species that thrive at both early and late successional stages can coexist. IDH is a nonequilibrium model used to describe the relationship between disturbance and species diversity. IDH is based on the following premises: First, ecological disturbances have major effects on species richness within the area of disturbance. Second, interspecific competition results in one species driving a competitor to extinction and becoming dominant in the ecosystem. Third, moderate ecological scale disturbances prevent interspecific competition.

Xerosere is a plant succession that is limited by water availability. It includes the different stages in a xerarch succession. Xerarch succession of ecological communities originated in extremely dry situation such as sand deserts, sand dunes, salt deserts, rock deserts etc. A xerosere may include lithoseres and psammoseres.

<span class="mw-page-title-main">Hydrosere</span>

A hydrosere is a plant succession which occurs in an area of fresh water such as in oxbow lakes and kettle lakes. In time, an area of open freshwater will naturally dry out, ultimately becoming woodland. During this change, a range of different landtypes such as swamp and marsh will succeed each other.

<span class="mw-page-title-main">Secondary succession</span>

Secondary succession is the secondary ecological succession of a plant's life. As opposed to the first, primary succession, secondary succession is a process started by an event that reduces an already established ecosystem to a smaller population of species, and as such secondary succession occurs on preexisting soil whereas primary succession usually occurs in a place lacking soil. Many factors can affect secondary succession, such as trophic interaction, initial composition, and competition-colonization trade-offs. The factors that control the increase in abundance of a species during succession may be determined mainly by seed production and dispersal, micro climate; landscape structure ; bulk density, pH, and soil texture.

<span class="mw-page-title-main">Disturbance (ecology)</span> Temporary change in environmental conditions that causes a pronounced change in an ecosystem

In ecology, a disturbance is a temporary change in environmental conditions that causes a pronounced change in an ecosystem. Disturbances often act quickly and with great effect, to alter the physical structure or arrangement of biotic and abiotic elements. A disturbance can also occur over a long period of time and can impact the biodiversity within an ecosystem.

"Auto-" meaning self or same, and "-genic" meaning producing or causing. Autogenic succession refers to ecological succession driven by biotic factors within an ecosystem and although the mechanisms of autogenic succession have long been debated, the role of living things in shaping the progression of succession was realized early on. Presently, there is more of a consensus that the mechanisms of facilitation, tolerance, and inhibition all contribute to autogenic succession. The concept of succession is most often associated with communities of vegetation and forests, though it is applicable to a broader range of ecosystems. In contrast, allogenic succession is driven by the abiotic components of the ecosystem.

<span class="mw-page-title-main">Plant ecology</span> The study of effect of the environment on the abundance and distribution of plants

Plant ecology is a subdiscipline of ecology that studies the distribution and abundance of plants, the effects of environmental factors upon the abundance of plants, and the interactions among plants and between plants and other organisms. Examples of these are the distribution of temperate deciduous forests in North America, the effects of drought or flooding upon plant survival, and competition among desert plants for water, or effects of herds of grazing animals upon the composition of grasslands.

<span class="mw-page-title-main">Climax species</span> Plant species that can germinate and grow with limited resources

Climax species, also called late seral, late-successional, K-selected or equilibrium species, are plant species that can germinate and grow with limited resources; e.g., they need heat exposure or low water availability. They are the species within forest succession that are more adapted to stable and predictable environments, and will remain essentially unchanged in terms of species composition for as long as a site remains undisturbed.

<span class="mw-page-title-main">Riparian-zone restoration</span> Ecological restoration of river banks and floodplains

Riparian-zone restoration is the ecological restoration of riparian-zonehabitats of streams, rivers, springs, lakes, floodplains, and other hydrologic ecologies. A riparian zone or riparian area is the interface between land and a river or stream. Riparian is also the proper nomenclature for one of the fifteen terrestrial biomes of the earth; the habitats of plant and animal communities along the margins and river banks are called riparian vegetation, characterized by Aquatic plants and animals that favor them. Riparian zones are significant in ecology, environmental management, and civil engineering because of their role in soil conservation, their habitat biodiversity, and the influence they have on fauna and aquatic ecosystems, including grassland, woodland, wetland or sub-surface features such as water tables. In some regions the terms riparian woodland, riparian forest, riparian buffer zone, or riparian strip are used to characterize a riparian zone.

The Janzen–Connell hypothesis is a widely accepted explanation for the maintenance of tree species biodiversity in tropical rainforests. It was published independently in the early 1970s by Daniel Janzen and Joseph Connell. According to their hypothesis, host-specific herbivores, pathogens, or other natural enemies make the areas near a parent tree inhospitable for the survival of seedlings. These natural enemies are referred to as 'distance-responsive predators' if they kill seeds or seedlings near the parent tree, or 'density-dependent predators' if they kill seeds or seedlings where they are most abundant. Such predators can prevent any one species from dominating the landscape, because if that species is too common, there will be few safe places for its seedlings to survive. However, because the predators are host-specific, they will not harm other tree species. As a result, if a species becomes very rare, then more predator-free areas will become available, giving that species' seedlings a competitive advantage. This negative feedback allows the tree species to coexist, and can be classified as a stabilizing mechanism.

Joseph Hurd Connell FAA was an American ecologist. He earned his MA degree in zoology at the University of California, Berkeley and his PhD at Glasgow University. Connell's first research paper examined the effects of interspecific competition and predation on populations of a barnacle species on the rocky shores of Scotland.</ref> According to Connell, this classic paper is often cited because it addressed ecological topics that previously had been given minor roles. Together, with a subsequent barnacle study on the influence of competition and desiccation, these two influential papers have laid the foundation for future research and the findings continue to have relevance to current ecology. His early work earned him a Guggenheim fellowship in 1962 and the George Mercer Award in 1963.

<i>Arctostaphylos rubra</i> Species of flowering plant

Arctostaphylos rubra is a species of flowering plant in the heath family and the genus Arctostaphylos, the manzanitas and bearberries. Common names include red fruit bearberry, alpine bearberry, arctic bearberry, red manzanita, and ravenberry. It is native to Eurasia and northern North America from Alaska through most of Canada to Greenland. There is also one population in the contiguous United States, located in the Absaroka Mountains of Wyoming.

<span class="mw-page-title-main">Gap dynamics</span>

Gap dynamics refers to the pattern of plant growth that occurs following the creation of a forest gap, a local area of natural disturbance that results in an opening in the canopy of a forest. Gap dynamics are a typical characteristic of both temperate and tropical forests and have a wide variety of causes and effects on forest life.

<span class="mw-page-title-main">UW Bothell Wetland Restoration Project</span>

The UW Bothell/Cascadia College Wetland Restoration Project is a 58-acre forested floodplain restoration site at the delta of North Creek in King County, Washington, USA. The State of Washington bought the site in 1994 from the Truly family, and dedicated the land to the construction of the Bothell regional campus of the University of Washington and Cascadia College. Construction began in 1998, as did the stream and floodplain restoration. Two years later, in 2000, classes opened for students, and in 2001 the main phase of the restoration was completed. Today the site is an ongoing area of restoration education.

References

  1. Ricklefs, 393-395.
  2. 1 2 Connell & Slatyer
  3. 1 2 3 4 5 6 7 8 9 Connell & Slatyer, 1121-1124.
  4. 1 2 Ricklefs, 400.
  5. Walker & Chapin.
  6. Walker & Chapin, 1518.
  7. Walker, et al.
  8. 1 2 3 Connell & Slatyer, 1124.
  9. "Sand Dunes".
  10. 1 2 Moorcroft.
  11. Wang.
  12. Connell & Slatyer, 1124-1127.
  13. Ricklefs, 401.
  14. Connell & Slatyer, 1125-1126.

Works cited