
Environmental Encyclopedia 3
Succession
As early successional species become established, they
alter their
environment
and make it more habitable for
later seral stages. Usually a disturbed or barren area has low
soil
nutrient
levels, intense sunlight, and no protection from
violent weather. Because precipitation quickly runs off the
bare ground, little moisture is available for plant growth.
Species that can survive under such harsh conditions have
little
competition
, and they spread quickly. As they grow
and thicken, these plants add organic matter to the soil,
which aids moisture retention and helps soil bacteria to grow.
As soil nutrients and moisture increase, larger shrubs and
perennial plants can take root. The shade of these larger
species weakens and eventually eliminates the original pion-
eering species, but it cools the local environment, further
improving moisture availability and allowing species that
demand relative environmental
stability
, such as woodland
species, to begin moving in. Eventually, shade-tolerant spe-
cies of plants will come to dominate the area that sun-loving
plants had first colonized.
One of the most well-documented examples of pri-
mary succession occurred in 1883 when the
volcano
on the
Indonesian island of
Krakatoa
erupted, destroying most of
the island and its life forms, and leaving a new island of
bare volcanic rock and ash. Within a few years several species
of grasses, ferns, and flowering shrubs had managed to arrive,
carried by wind, water, or passing birds from islands 25 mi
(40 km) or more distant. Just 40 years later, more than 300
plant species were growing, and in some areas over 12 in
(30 cm) of soil had developed: soil bacteria, insects, and
decomposers
had managed to reach the island and were
turning fallen organic
detritus
into soil. Over the decades,
the total number of species gradually stabilized, but the
character of the community continued to change as incoming
species replaced earlier arrivals. On Krakatoa succession hap-
pened with unusual speed because the
climate
is warm and
humid and because the fresh volcanic ash made a nutrient-
rich soil.
Early successional species, those most able to quickly
establish a foothold, are known as pioneer species. Usually
pioneer species are opportunists, able to find nourishment
and survive under a great variety of conditions, quick to grow,
and able to produce a great number of seeds or offspring at
once. Pioneer species have very effective means of dispersing
seeds or young, and their seeds can often remain dormant
in soil for some time, sprouting only after conditions become
suitable. Dandelions are well-known pioneers because they
can quickly produce a seed head with hundreds of seeds.
Each tiny seed has a lightweight structure that allows the
wind to carry it long distances. Dandelions are quick to
invade a lawn because of this effective seed-dispersal tactic,
because they can survive under sunny or shady conditions,
and because they grow and reproduce quickly. In open sun-
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shine, they are competitive enough to establish themselves
despite the presence of a thick mat of turf grass.
Later successional species tend to be more shade toler-
ant, slower to grow and reproduce, and longer lived than
pioneer species. Their larger seeds do not disperse as easily
(compare the size of an acorn or walnut with that of a
dandelion seed), and their seeds cannot remain dormant for
very long before they lose viability. Seedlings of some late
successional species, such as the Pacific Northwest hemlock,
require shade to survive, and most require considerable mois-
ture and soil nutrients.
We usually think of succession occurring after a cata-
strophic environmental disturbance, but in some cases the
gradual environmental changes of succession proceed in the
absence of disturbance. Two outstanding examples of this
are bog succession and the invasion of prairies by shrubs
and trees.
In cool, moist, temperate climates, plant succession
often turns ponds into forest through bog succession. Water-
loving plants, mainly rush-like sedges and sphagnum moss,
gradually creep out from the pond’s edges. Often these plants
form a floating mat of living and dead vegetation. Other
plants—cranberries, Labrador tea, bog rosemary—become
established on top of the mat. Organic detritus accumulates
below the mat. Eventually the bog becomes firm enough to
support black spruce, tamarack, and other tree species. As
it fills in and dries, the former pond slowly becomes indistin-
guishable from the surrounding forest.
Many
grasslands
persist only in the presence of occa-
sional wildfires. During wet decades or when human activity
prevents fires, woody species tend to creep in from the edges
of a
prairie
. If sufficient moisture is available and if fires
do not return, open grassland can give way to forest. Fire
suppression has aided the advance of forests in this way
across much of the United States, Canada, and Mexico.
Grazing or browsing animals can also be important forces
in maintaining biological communities. Adding or removing
grazers can initiate successional processes.
Longer, slower disturbances than fire or field clearing
can also initiate succession. Over the course of centuries,
climate change can cause significant alteration in the charac-
ter of biomes. Minor variations in rainfall or temperature
ranges can alter community structure for decades or centu-
ries. The end of the last glacial period about 10,000 years
ago allowed
tundra
, then grasslands, then temperate forests
to advance northward across North America. Even geologic
activity, such as mountain building or changes in sea level,
has caused succession. More recently, human introductions
of species from one continent to another have caused signifi-
cant restructuring of some biological communities.
Turnover from one community to another may occur
in just a few years or decades, as it did on Krakatoa. Stages