Succession and Stability

In 1794, Captain George Vancouver visited the inlet to what is today called Glacier Bay, Alaska
pass. He could not beyond the inlet to the bay, however, because his way was blocked by a
mountain of ice. Vancouver (1798) described the scene as follows: "The shores of the continent
form two large open bays which were terminated by compact solid mountains of ice, rising
perpendicularly from the water's edge, and bounded to the north by a continuation of the united
lofty frozen mountains that extend eastward from Mount Fairweather."
In 1879, John Muir explored the coast of Alaska, relying heavily on Vancouver's earlier
descriptions. Muir (1915 ) commented in his journal that Vancouver's descriptions were excellent
guides except for the area within Glacier Bay. Where Vancouver had met "mountains of ice," Muir
found open water. He and his guides from the Hoona tribe paddled their canoe through Glacier
Bay in rain and mist, feeling their way through uncharted territory. They eventually found the
glaciers, which Muir estimated had retreated 30 to 40 km up the glacial valley since Vancouver's
visit 85 years earlier.
Muir found no forests at the upper portions of the bay. He and his party had to build their
campfires with the stumps and trunks of long-dead trees exposed by the retreating glaciers. Muir
recognized that this "fossil wood" was a remnant of a forest that had been covered by advancing
glaciers centuries earlier. He also saw that plants quickly colonized the areas uncovered by
glaciers and that the oldest exposed areas, where Vancouver had met his mountains of ice,
already supported forests.
Muir's observations in Glacier Bay were published in 1915 and read the same year by the
ecologist William S. Cooper Encouraged by Muir's descriptions, Cooper visited Glacier Bay in
1916 in what was the beginning of a lifetime of study. Cooper saw Glacier Bay as the ideal
laboratory for the study of ecological succession, the gradual change in plant and animal
communities in an area following disturbance or the creation of flew substrate. Glacier Bay was
ideal for the study of succession because the history of glacial retreat could be accurately traced
back to 1794 and perhaps farther.
Cooper ultimately made four expeditions to Glacier, Bay. His work and that of later ecologists
produced a detailed picture of succession there. Several species of plants colonize an area during
the first 20 years after it is expose by the retreating glacier. These plants, the first in a
successional sequence, form a pioneer community. The most common members of the pioneer
community are horsetail, Equisetum varietaum, willow herb, Epilobium latifolium, willows, Salix
sp., cottonwood seedlings, Populus balsamifera, mountain avens, Dryas drurnmondii, and Sitka
spruce, Picea sitchensis.
About 30 years after an area is exposed, the pioneer community gradually grades into a
community dominated by mats of Dryas, a dwarf shrub. These Dryas mats also contain scattered
alder, Alnus crispa, Salix, Populus, and Picea. Then, about 40 years after glacial retreat, the
community changes into a shrub-thicket dominated by Alnus. Soon after the closure of the Alnus
thicket, however, Populus and Picea will grow above it, covering about 50% of the area on sites
50 to 70 years old.
In 75 to I00 years, succession leads to a forest community dominated by Picea. Mosses carpet
the understory of this spruce forest and hem and there grow seedlings of western hemlock,
Tsuga heterophylla, and mountain hemlock, Tsuga mertensiana. Eventually, the population of
Picea declines and the forests are dominated by Tsuga. On landscapes with shallow slopes these

hemlock forests eventually give way to muskeg, a landscape of peat bogs and scattered tussock
meadows.
Because succession around Glacier Bay occurs on newly exposed geological substrates, not
significantly modified by organisms, ecologists refer to this process as primary succession,
Primary succession also occurs on newly formed volcanic surfaces such as lava flows. In areas
where disturbance destroys a community without destroying the soil, the subsequent succession
is called secondary succession. For instance, secondary succession occurs after agricultural lands
are abandoned or after a forest fire.
Succession generally ends with a community whose populations remain stable until disrupted by
disturbance. This late successional community is called the climax community. The nature of the
climax community depends upon environmental circumstances. The communities we discussed
in chapter 2temperate forests, grasslands, etc.were essentially the climax communities for
each of the climatic regimes that we considered. The climax community around Glacier Bay is
determined by the prevailing climate and local topography. On well-drained, steep slopes the
climax community is hemlock forest. In poorly drained soil on shallow slopes the climax
community is muskeg.
Studies of succession show that communities and ecosystems are not static but constantly
change in response to disturbance, environmental change, and their own internal dynamics. In
many cases, the general direction of change in community structure and ecosystem processes is
predictable, at least over the short term. The patterns of change in community and ecosystem
properties during succession and the mechanisms responsible for those changes are subjects
covered in this chapter. We also consider a companion topic, community and ecosystem stability.
CONCEPTS

Community changes during succession include increases in species diversity and changes
in species composition.
Ecosystem changes during succession include increases in biomass, primary production,
respiration, and nutrient retention.
Mechanisms that drive ecological succession include facilitation, tolerance, and inhibition.
Community stability may be due to lack of disturbance or community resistance or
resilience in the face of disturbance.

Some of the most detailed studies of ecological succession have focused on succession leading
to a forest climax. Though primary and secondary forest succession require different amounts of
time, the changes in species diversity that occur in each appear remarkably similar. Primary
Succession
At Glacier Bay We have already reviewed the basic patterns of primary succession around Glacier
Bay. Now we return to Glacier Bay to examine successional changes in species diversity and
composition. William Reiners, Ian Worley, and Donald Lawrence (1971) studied changes in plant
diversity during succession at Glacier Bay. They worked at sites carefully chosen for similarity in
physical features but differing substantially in age. Their eight study sites were below 100 m
elevation, were on glacial till, an unstratified and unsorted material deposited by a glacier, and
all had moderate slopes. The study sites ranged in age, that is, time since glacial retreat, from 10
to 1,500 years.

Their youngest site, which was approximately 10 years old, supported a pioneer community of
scattered Epilobium, Equisetum, and Salix. Site 2 was about 23 years old and supported a mix of
pioneer species and clumps of Populus and Dryas. Site 3, which was approximately 33 years old,
supported a mat of Dryas enclosing clumps of Salix, Populus, and Alnus. Site 4 was 44 years old
and was dominated by a mat of Dryas with few open patches. Site 5, which was approximately
108 years old, was dominated by a thicket of AInus and Salix with enough emergent Populus and
Picea to form a parrial canopy. Site 6 was a 200-year-old forest of Picea. Using geological
methods, Reiners and his colleagues dated site 7 at 500 years and site 8 at 1,500 years. Both
sites were located on Pleasant Island, which because it is located outside the mouth of Glacier
Bay had escaped the most recent glaciation, which had destroyed the forests along the bay. Site
7 was an old forest of Tsuga that contained a few Picea. Site 8 was a muskeg with scattered
lodgepole pines, Pinus contorta.
The total number of plant species in the eight study sites increased with plot age. As you can see
in figure 20.2, species richness increased rapidly in the early years of succession at Glacier Bay
and then more slowly during the later stages, approaching a possible plateau in species richness.

FIGURE 20.2 Change in plant species richness during primary succession at Glacier Bay, Alaska
(data from Reiners. Worley, and Lawrence 1971).
Not ail groups of plants increased in diversity throughout succession. Figure 20.3 shows that
while the species richness of mosses, liverworts, and lichens reached a plateau after about a
century of succession, the diversity of low shrubs and herbs continued to increase throughout
succession. In contrast, the diversity of tall shrubs and trees increased until the middle stages of
succession and then declined in later stages.

FIGURE 20.3 Succession of plant growth forms at Glacier Bay, Alaska (data from Reiners, Worley,
and Lawrence 1971).
The pattern of increased species richness withstand age that Reiners and his colleagues
described for the successional sequence around Glacier Bay is one that we will see several times
in the examples that follow. However, the tempo of succession is far different. The late
successional climax community at Glacier Bay was 1,500 years old. In the following example of
secondary succession, the climax forest community was 150 to 200 years old, approximately
one-tenth the age of the climax community studied at Glacier Bay.
Secondary Succession in Temperate Forests
The Piedmont Plateau of eastern North America includes some of the most convenient places to
study secondary succession. The deciduous forests of this region were intensively cleared and
cultivated beginning approximately three centuries ago. As fields new forest areas cleared, the
region was progressively converted to a patchwork of abandoned fields of ions ages interspersed
with a few areas of undisturbed forest.
This situation provided Henry J. Oosting (1942) with study sites in virtually every stage of
secondary succession for his now classic studies of succession in the Piedmont Plateau of North
Carolina. David Johnston and Eugene Odum (1956) described the pattern of succession on the
Piedmont Plateau as follows. The first species to colonize and dominate abandoned fields are
crabgrass, Digitaria sanguinalis, and horseweed, Erigeron canadense. During the second year of
succession, the fields are dominated by either aster, Aster pilosis, or ragweed, Ambrosia
artemisiifolia. A few years later the field is covered by broomsedge, Andropogon virginicus, with
a scattering of other shrubs and small trees. Pine seedlings may appear as early as the third year
and may form a closed canopy in 10 to 15 years. Pine seedlings cannot grow in the shade of
larger pines but the seedlings of many deciduous trees can. Consequently, 40- to 50-year-old
pine forests generally have a well-developed understory of young deciduous trees. These
deciduous trees, especially oak, Quercus, and hickory, Carya, become the dominant trees by
about 150 years, and the pines decline to a few scattered individuals. Because Quercus and
Carya can reproduce in their own shade, the late successional oak-hickory forest is considered
the climax stage.
Oosting's data show that the number of woody plant species increased during secondary
succession on the Piedmont Plateau (fig. 20.4). The successional sequence began with a single

species of woody plant invading soon after fields were abandoned and began to level off at 50 to
60 species after approximately 150 years. This increase in species richness follows a logarithmic
pattern like the one we saw' at Glacier Bay.

FIGURE 20.4 Change in woody plant species richness during secondary forest succession in
eastern North America (data from Oosting 1942).
How does animal diversity change across the same successional sequence? Johnston and Odum
studied the birds living on thirteen 20-acm study sites ranging in successional age from 1 to 150
years and supporting vegetation ranging from grassland to mature oak-hickory forest. The
increase in bird diversity across this successional sequence closely paralleled the increase in
woody plant diversity observed by Oosting (fig. 20.5). Duringthe grass-forb stage of succession
the bird community consisted of two species generally associated with grasslands. In the grassshrub stage, the diversity of birds increased to 8 to 13 species. In 25- to 35-year-old pine forests,
the diversity of birds was 10 to 12 species. Pine forests of 60 to 100 years supported 23 to 24
bird species. Johnston and Odum recorded 22 bird species in old-growth oak-hickory forests.

FIGURE 20.5 Change in number of breeding bird species during secondary forest succession
(data from Johnston and Odum I956).

In summary, over a period of approximately one and a half centuries abandoned fields in eastern
North America undergo successional changes in plant and bird communities that involve changes
in species composition and increases in species diversity. Similar successional changes occur in
the marine intertidal zone but on an even shorter timescale: approximately one and a half years
instead of one and a half centuries.