In terms of Ecology, the term ‘succession’ can be defined as ‘The change in species composition, community structure and function over time’ (MacKenzie et al 2001).
Succession of plant species follows a disturbance or initial colonisation of a new habitat. Take a back garden: the owners abandoned the house, stopped mowing the lawn, and the garden was left completely to its own devices. Eventually, after many stages of development, this garden would probably become a forest.
Grass is a competitive species, however with no regular mowing taking place; new opportunistic pioneering species (r-selected species) which grow easily quickly begin to take hold. Then, shrubs begin to jump in and smaller species die out. Eventually, tree saplings (K-selected species) grow large enough to dominate, and compete with smaller species by engulfing all the light, water and nutrients. The final set of ruling species is called the ‘climax community’.
Take a look around and begin to imagine what could be under your feet if nature was to take its course. The grass fields all around us are what is called a ‘plagioclimax’– this is, a climax community caused by human halting of natural succession. Should these fields be forest? Was the Oak woodland always like that or perhaps planted by humans? How about the dandelions that grow between the cracks of tarmac at a dis-used industrial site; are they, perhaps the first in the line of succession that would eventually become a natural jungle?
Primary succession; succession that occurs on newly formed surfaces (with no seed bank) such as dunes, lava flows and glacial forelands takes hundreds or thousands of years to facilitate. Secondary succession; succession that begins on previously vegetated areas such as fallow fields, clear-cut forests and embankments, vegetates rapidly.
Succession takes two main forms. ‘Autogenic’: natural, plant driven succession uninhibited by outside abiotic factors; and ‘Allogenic’: succession driven by abiotic, environmental factors such as human intervention, grazing animals, water flows, etc. Allogenic succession is far more common- true autogenic succession requires perfect growing conditions for all species, and virtually no human interference.
Henry Cowels (1899) was one of the first scientists to document this observable phenomenon. Using the example of his study site, Lake Michigan Sand Dunes, he noted the succession of species. Succession began with Marram Grass, which would facilitate within 20 years. Cotton Wood proceeded after 20-50 years, Jack Pine after 50-100 years, and Black Oak eventually dominates after 100-150 years. Each of these stages is commonly referred to as a ‘sere’.
In 1916 Fredrick Clements named this ‘facilitation’: complete replacement of species, where the preceding species inadvertently makes conditions habitable, or more favourable to other species. Clements held that ‘after a disturbance, an ecosystem would eventually return to its characteristic species assemblage’. He believed that succession was somewhat predicable and that eventually one would observe a monoclimax– the ‘King’ species of succession.
However, ‘many studies have shown successional pathways to be much more complicated and unpredictable than the classic facilitation model’ (Mac Donald 2003).
In the early 1920’s, Henry Gleason offered a more complex alternative to Clements’ model, suggesting that species responded individually to environmental factors and that the community structure was less predictable and more ‘coincidental’.
Years after Cowels’ benchmark study of Lake Michigan Sand Dunes, Jerry Olson (1958) conducted a long term study at the same site and concluded that the final, ‘climax’ community or assemblage differed depending on original conditions. He developed the idea of ‘multiple successional pathways’, depending on original conditions, extrinsic variables and colonisation patterns.
However, apart from environmental conditions and external factors, during the latter half of the 20th century, the more complex relationships between plant species that would eventually begin to explain the differing colonisation patterns were being researched.
In 1950, ecologist Catherine Keever observed succession following field abandonment and detected allelopathic and competitive interactions taking place. Allelopathy is a biological phenomenon where one species releases a biochemical which hinders growth, survival and/or reproduction of another species. Keever writes that Broomsage dominates as the ‘King’ species for this reason.
It soon became apparent that there is no simple pattern of succession. Instead, an array of intertwining mutualistic and competitive relationships built upon a combination of existing conditions, external factors and species interrelationships. This means, similarly to a food web among the animal kingdom, a change in one factor or species can alter the eventual climax community.
In 1977 Joseph Connell and Ralph Slatyer published their work which summarised the three basic modes in which an ecological community interact. The Conell-Slatyer model of succession includes three broad mechanisms: Facilitation, Inhibition and Tolerance.
Facilitation is ‘competitive replacement of species’. Each community creates conditions favourable for more complex communities, and are eventually ‘overtaken’ by more competitive species. Tolerance is where all species involved are equally capable of establishing themselves, and tolerate conditions put upon them by other species (such as lack of light) in order to survive. Late successional species are incredibly tolerant and competitive. Lastly, Inhibition is where initial plants modify the environment so that it is less favourable to other species, such as through nutrient sapping and allelopathy.
Lawrence Walker and colleagues observed all three models taking place at once during a 7 year study in Puerto Rico. During the study, early successional forms and woody plants were removed, which eventually affected the long term successional species- it appeared that woody plants and ferns appeared to, in combination, facilitate long term forest development (Walker et al 2001).
In contrast, scrambling ferns inhibited succession and decreased woody plant richness. This phenomenon can be attributed to a host of different and interlinking factors such as alteration of soil pH and light availably.
Walker’s study highlights problems such as species arrival order, which make projections almost impossible when studying the contrasting effects of competition and facilitation. It becomes clear that, ‘although it is an observable site phenomenon, there is no unanimity of definition or casual explanation for the process’ (Joy Tivvy, 1993).
So if you go down to the woods today, you’re almost sure of a big surprise!