Community Ecology II: Ecological Successions

I. Introduction

A. Old Classical Field of Ecology

1. Early observation of the predictability of ecological successions by Kerner

(1863) in Danube Basin and, Hult (1885) in Finland, as summarized by
Warming (1895), were important in the development of ecology as a science.

2. The study of successions became a dominant theme in plant ecology

especially in North America because of the pattern of exploitative agriculture
where farmers moved on after wearing out the soil of their farms.
Consequently patterns of ecological successions were everywhere to be
studied.
3. While most of the discussion here emphasizes terrestrial plant ecology,
successions occur every where, as shown in Smiths Figs. 21.2, 21.6, 21.11-
21.13, 21.18.

B. Clementsian School

1. Started with the work of Cowles (1899) of the University of Chicago on Lake
Michigan at what is now called the Indiana Dunes Park and National Sea Shore.
At this location are an interesting series of dunes and ponds.

a. Cowles discovered that the dunes and ponds closest to Lake Michigan
were the youngest.

b. The dunes and pond furthest away form the Lake were the oldest.

c. The ecological succession starting with the ponds filling in, going from
oligotrophic to eutrophic to wetland marshes to swamps, to wet forest
and then to the Beech-maple climax characteristic of the area.

d. At the same time the dunes were stabilized by plants, first grasses, then
shrubs, then Oak-Hickory, and finally the Beech-Maple Climax
community.

e. This means different physical starting points converged on the same

climax Beech-Maple Forest.

2. Clements used Cowles's work as the basis for his organismal/ superorganism
theory of the community — Communities develop like organisms. Plant
Successions (1916) was written by Clements in support of his holistic view.
 3. This research was largely American, in reflection of the pioneer exploration
and recovery of farmland by forest as farmers pushed westward.

4. Gleason (1926) was very skeptical of this view as was discussed earlier.

II. Classical terminology and concepts

A. Climax — was defined by Clements as an enduring natural community in

equilibrium with climate.

1. Integrated with the superorganism paradigm of Clements — he developed the

idea as a fundamental ecological process.

2. This concept of the climax was based on long-term adaptations of

communities to climate (Clements 1936, Nature and Structure of the Climax).

3. As we earlier discussed, Whittaker proposed that the climax should be

operationally defined as self-sustaining enduring end point of succession.

B. Sere — A Complete Ecological Succession

1. Pioneer earliest invaders —> seral stages — >climax. This is a sere.

2. Ecesis is successful colonization, growth and reproduction by a species in a

succession.

3. Associes are temporary successional communities as defined by Clements.

4. Subclimax is a long lasting, usually a forest successional seral stage.

5. Convergence of xerarch seres and hydrarch seres in the Indiana dunes area led
to same climatic climax, which led Clements to believe this validated his
superorganism concept of the climax as being the community adapted to the
climate.

6. The succession started with a disturbance that results in a nudation, the

elimination of the original community.

C. Primary Succession (See Smiths Figs. 21.6-21.9)

1. Natural bare areas, rock, lava, sand, water—complete nudation.

2. Slow succession occurring through physical and biological processes.

.
a. Physical changes — Allogenic changes — create soil profile.
 b. Biological changes — Autogenic changes — changes in biological
community.

D. Secondary Successions

1. Climax vegetation destroyed by man or nature — fire, hurricanes, etc. but

more usually in the USA through cutting of the forest for agriculture.

2. Because the soil was already there, there is a rapid return to climax from
fallow field, mostly by autogenic, not allogenic changes.

3. Fugitive species have a hit and run adaptation to rapid dispersal and ecesis
especially adapted to the early succession stages, but cannot compete later in the
succession.

4. Billings (1938) and Keever (1950) in North Carolina carefully studied the
secondary successions and came up with the following pattern of succession
common in the southern coast plain of the USA.

Crab grass, weeds (one year)Æ Horseweed field (two years)Æ Aster Composite
stage (3 yrs).Æ sedges (5 to 15 yrs)Æ Pine forest (50 to 150 yrs) Æ Oaks.

E. Modifiers of climax

1. Sereclimax — long-persisting successional stage due to some interrupting or

slowing fate -- like poor soil or pine barrens stuck in succession to Clements
these are merely taking a long time to provide the allogenic changes that allow
the climatic climax to establish itself.

2. Subclimax — long-persisting forest stage before final forest climax.

3. Disclimax — A community prevented from going to climax by an interfering

factor like fire, overgrazing, i.e. a “bad” apparent climax.

F. Climax reconsidered

1. Monoclimax adapted to the climate was Clements’s concept. This is not far
from our earlier discussions of the relationship in terrestrial ecology between
the formations, climate and the soil. His view was extremely long in
perspective, he saw mountains being eroded down, and many communities as in
mountains were only associations on their way to the ultimate climatic climax.

2. Some ecoogists as Tansley conceived of the Polyclimax — that there were

many other possibilities for the climax based on other physical conditions than
climate, and of course, as we discussed earlier, Gleason threw the climax idea
out along with the superorganism, thinking that all there were are individual
organisms, all else were imaginary, even the polyclimax.
3. Climax pattern theory: ecoclines: continua of Whittaker. As discussed,
Whittaker saw the world in terms of varying environmental parameters, and
that species were adapted to conditions along the gradients. However, some
species that are dominants determine, as a keystone species, the ecology of a
community, so these species have great importance. Whittaker then defines the
climax as a community that endures and replaces itself.

4. Whittaker would say then, that fire as a natural factor, can determine the
climax for many communities: All the grassland formations, the chaparral, and
many coniferous forests, such as the Piney Woods of East Texas and the Gulf
coast require fire and are therefore pyroclimaxes. Some pines in the west,
even have cones that open only after a fire. Often one sees a forest of the same
age trees that all started after a fire.

5. Grazing in grasslands is similarly tolerated by grasses for the same reason, as

grasses grow from their bases not the apex. Thus fire and grazing work together
to maintain the grassland biome.

6. Bormann and Likens noted that because trees eventually die and fall,
creating an opening and opportunity for other plants to enter, proposed that
many, if not all climaxes, are actually better called shifting mosaic steady state
climaxes. In Maple-Beech Forest, the dominant climax community in much of
northeastern North America, maple seedlings tend to grow under Beeches and
the beech seedlings tend to grow under Maples, a reciprocal replacement.

7. Extreme age an important strategy -– very few old growth forests left —
Douglas fir 450-1000 years old — is replaced by Western Hemlock and silver
fir–enormous trees with huge biomass. A unique fauna including the spotted
owl, tree vole, etc. have endured to live here in this special ancient forest.

8. Even Sequoia sempervirens, the coast redwood, and Sequoiadendron

giganteum, the Big Tree, are fire climax forests and require open sunny areas
for the regeneration of their seedlings, and use their great longevity to produce
seeds over a long time toward the time that conditions will be advantageous for
their seedlings to survive in the sun after a fire. Here age and fire resistance
operate together as a strategy for existence.

9. Edaphic climaxes are based on soil influences. The "Lost Pines" of Bastrop
County are a good example. They are adapted to sandy soils, while the grasses
are adapted to clayey soils of the black land prairies--both are fire climaxes.

10. Plagioclimaxes are communities maintained by man and his grazing

animals. Many parts of the world have been used--or misused by man for so
long that we do not know what is .the original or expected climax vegetation.
 11. Desertification — reverses primary succession — loss of water holding
ability of the soil changes the climax through human interference — this is a
very serious problem. Often the ‘plagioclimax” is the consequence of
desertification. Roman roads leading out into the desert in North Africa give
testimony to the destructive effect of Mankind. North Africa was the granary of
Rome.

12. Climatic Change—Changes in climate since the Pleistocene and ongoing

changes causes changes in the prevailing climax as revealed by palynology and
studies now show the advance of warm-adapted species and the retreat
northward of cold-adapted species in probable response to global warming.

G. Succession causes as suggested by Egler (1954)

1. Ecesis: establishment and colonization of propagates in the succession is basic

to the progressive changes in organisms present in the successions.

2. Relay Floristic (and faunistics) — one seral stage gives way to another
progressively, like a relay race each species passing on the baton. This is based
on the plants modifying the environment allogenically, allowing later
successional plants to come in and replace the earlier stages. This corresponds
to Primary Successions.

3. Initial Floristic (and faunistics) Composition—Here all the plants start out
together in terms of propagules— firstest with mostest starts the succession, and
the later plants come in and replace the earlier plants by competition. This is
basically the secondary succession of Clements.

H. Succession Models of Connell and Slatyer (1977)

1. Facilitation model: the autogenic changes brought about by the earlier stages
help later successional plants to take over, and is similar to the floristic relay.

2. Inhibition model: The first invaders render the community less suitable for
latter succession, slows succession by inhibition, but the longer lived plants out
compete eventually the early succession plants. This is somewhat similar to the
floristic composition mode.

3. Tolerance model: Here the plants that are present have no effect on later
succession plants, there is no facilitation or inhibition as such, the most tolerant
species win out in this competition.

4. All of these theories involve changes, allogenic and autogenic, and all are
restatements in various ways of the original Clementsian ideas to make them
more reductionist, less holistic in the sense of Clements’s superorganism.
 5. While much of the emphasis is on plants as they create the biomass of the
community in terrestrial systems, successions greatly affect the animals, too,
as seen in Smiths Fig. 21.19, where different animal species live in different
stages of the succession. Many species are edge species. Others are interior
species and only live in the old growth forest. Others require both.

6. Replacement in successions: Disturbances

Organisms have life spans, thus constant, continuous replacement occurs as in
a forest canopy, the shifting mosaic patterns of the climax forests as
emphasized by Bormann and Likens.

7. Cyclic Succession

a. Tundra ÅÆ Taiga: trees shade the soil, permafrost rises, trees fall over
through lack of soil for roots (the drunken forest); the tundra replaces it,
lets sunlight in to melt the permafrost, and again the trees enter.

b. Cycles of intermittent ponds and old-field moss and lichens show this
cycling of communities (see Fig. 21.14 in Smiths).

8. Some ecosystems are maintained by continual pulselike disturbances such as

the intertidal environments where the tides and storms continually disturb the
coastal environments (Figs. 21.11-21.13 in Smiths), or in wet-dry climates
where a forest in the dry season becomes a varzea swamp during the rainy
season.

9. Disturbances Æ perturbations, that is temporary changes in the biota.

a. Fires led to special fire-adapted communities that can handle the fire
disturbances with the least perturbations, especially of surface fires.

b. Hurricanes can lay level very large areas of forest.

c. Flooding can kill large areas of forest and promote special flood
forests. Animals (and man) can dam streams and flood the riparian
vegetation, causing it to change wetland vegetation.

d. Severe drought can cause great damage killing vegetation and

promoting fire in forests that do not normally burn.

e. Defoliating insects can cause the dead of large areas of forest by

exhausting the energy reserves of the plants.

f. Very large animals such as elephants in Africa can destroy woodland

by literally uprooting the trees allowing grass to spread, which then
maintains itself through fire and grazing (see Smiths Fig. 23.23-23.24).
 g. Man’s activities, agriculture, mining, lumbering can cause great
disturbances.

h. The repair of these perturbations occurs through ecological successions

small and large is a natural process.

10. Restoration Ecology is a branch of applied ecology that seeks to help an

ecosystem recover from great perturbation, usually those caused by man. In
the cases of plagioclimaxes, this can be a difficult task, and is often the first
step in trying to save endangered species, as it is often the very ecosystem that
is endangered.

11. Successions are an important part of Landscape Ecology, which studies how
ecosystems interact and how man affects landscapes, and much of the affects
are through disturbing natural communities, establishing plagioclimaxes,
agroecosystems, fragmenting communities, creating ecotones and patches.
(See Discussion in Smiths Chapter 23)

12. Fluctuations in Nature can be cyclic, bringing about regular perturbations in

the communities.

a. Water table changes can bring about changes because plants differ in
their ability to tolerate water-saturated soil. This can change with rise
and fall of the water table with changes in the rainfall.

b. Climatic shifts appear to be normal so the idea of ancient communities

that have existed since the Cretaceous are probably false and
communities even in the tropics have always had to adapt to such
changes and disturbances, although the climatic ones tend to very slow
acting, allowing time for communities to either adapt or migrate.

13. Frequency of disturbances important.

a. Occasional small ones are easier to handle, and are less perturbing
than ones that happen so frequently as to prevent the restoration of the
community through the healing of an ecological succession.
b. This is especially true of forest fires in the west where frequent small
fires are much less destructive than large fires (See Smiths Figs.
23.17-21.18).

J. Stability

1. Local — small perturbations in the community reflecting changes basically

local such as a small fire, or a flood along a stream. Ecological Successions
heal these quickly.
 2. Global — large-scale disturbances that bring about very large perturbations,
from which it may be difficult to restore community equilibrium. Some of the
disturbances of man can fall in this category.

3. Resistance: measures the amount of perturbation resulting from a disturbance.

a. The community with resistance has the resources to withstand the

disturbance and maintain itself.

b. This is characteristic of communities adapted to relatively non-stressful

environments in which disturbances are minor, of local impact.

c. However, such communities are often highly fragile, and cannot handle
major or global disturbance. Remember communities are the
consequence of natural selection on the constituent species, and
disturbance by man can be so great as to overwhelm the existing
resistance, shattering the ecosystem.

4. Resilience: is the ability to return to original conditions.

a. Although the perturbation may be great, the community bounces back

readily, even when the resistance is low as in many aquatic
communities.

b. Temperate communities normally encounter large disturbances, and are

consequently characterized by great resilience. Natural selection on the
constituent species would be for such adaptations.

c. That translates into resilience against even great perturbations caused by

man in temperate environments, contrasting with the fragility of most
tropical communities.

III. Processes and trends of successions (See Odum on page 23 in class notes) Smith unduly
casts doubt on several of the generalizations of Odum, which are based primarily on
secondary successions.

A. Species turnover basically goes from r-selected species to K-selected species

1. Rapid ecesis Æ slow ecesis. New communities exploiting the pioneer

situation need to establish quickly for the environment can be very short lived in
secondary successions

2. High dispersal capabilities ––> low dispersal capabilities. Usually the

nudations (places devoid of organisms) are scattered here and there, and to get
to them takes considerable ability for dispersal, while the climax community is
widespread, often everywhere, so the only need to disperse is for finding a host
plants insects, or to
 3. Shade intolerance — >Shade tolerance. Larger dominant plant species
overtop early successional plants. A vertical stratification of species occurs in
the forest with reference to tolerance of low light intensity

4. Short-lived plants—>long-lived plants, which are dominants that overgrow

the early successional plants, which are usually shade intolerant

5. Small seeds — >large seeds. The more endosperm present, the more energy
available to lift leaves to the sunlight and roots to grow into the ground

6. Low competition —>high competition. Early successional organisms are r-

selected in adaptation to a temporary environment. Late successional climax
communities are there for the long haul and are K-selected for survival so
competition is great

7. Defense mechanisms against grazers and browsers of plants increase from

early to late successional stages; biochemical diversity increases

8. Animal grazers change from low host specificity to high specificity in

response to the poisonous defense mechanisms of plants of the climax.

B. Species composition and interactions

1. Increase in diversity overall occurs through the succession although this is

obscured by the large size of the trees in the climax and the very small size of
the consumers, insects and the microbial life, the fungi and bacteria that attack
detritus and the tiny size of the feeders on microbial life.

2. Dominance becomes shared by many species. This is especially true of

tropical forests and the chaparral; in the temperate, especially boreal forest,
number of tree species is usually low in the climax.

3. Ecological niches become more specific: long-term species associations

develop in the climax.

4. More host plants, more places to live in vertical stratification of the climax as
in the tropical rain forests leads to greater species richness.

5. Symbioses become more richly developed: more time to develop relationships

such as mycorrhizae in forest plants. The larger biomass gives more places for
other organisms.

6. Species abundances stable — more resistant to change in the climax as the

species are mostly controlled by density-dependent factors of K-selection.

C. Energy and nutrients

 1. In early stages community primary productivity P (GPP) > Community R Æ
climax P = R of the whole community.

2. High maintenance costs on all that biomass: higher gross PP, less net PP
available to the higher trophic levels. Moreover, plants use energy resources to
create their defenses against herbivore grazers.

3. Shift from grazing to detritus food chains because animals restricted by

cellulose and lignins in plants as they become progressively larger and more
woody through the protection and support of the biomass mentioned above.

4. Detritus as humus modifies the soil, improving it, which is an important factor
in succession, in promoting the ecesis of later, larger plants.

5. Early successions have open nutrient cycling—> closed nutrient cycling, that
is, from nutrients in the soil to nutrients released from the biomass of detritus.

6. Little nutrients stored in the low biomass of the early succession to having
most of the nutrients locked up in biomass of the forest.

7. Development of symbiosis to enhance uptake and recycling of nutrients.

8. Results in a very low output or loss of nutrients from the climax community,
especially of tropical rainforests.

9. Organisms are adapted for the long term in terms of energy and nutrients;
while pioneers short-term rapid competition for quick growth and a place in
sun.

D. Man and succession

1. Consider that in agroecosystems net production goes to human consumption.

2. There is high net productivity in early successional stages.

a. This means there is rapid growth in biomass as maintenance costs are

low because the biomass of plants is small in the early succession.

b. Rapid net productivity, little investment in — defense mechanisms,

cellulose, lignin, and nutrient conservation.

c. Reliance on nutrients already easily available an open nutrient cycling

makes early successional plants amenable to using fertilizer.

3. Humans eat soft herbs and nonpoisonous seeds.

 4. Both are characteristic of plants of early successional stages; often the food
plants are closely related to the weeds in the garden.

5. Humans have genetically tailored the crop plants.

a. To pour energy into net of seeds and soft tissues that can we eat

b. Also this tends to reduce the plant’s ability to compete and protect itself,
leading to plants that cannot grow in nature, and we must protect our
plants from the herbivores as we have bred out of them resistance so we
can more readily eat them.

6. Even in forest plantations — subclimax of Piney Woods more economical crop

— few species, rapid growth, good harvest of lumber compared to the species-
rich hardwood climax of angiospermous trees.

7. Man must therefore invest much energy, an energy subsidy of fossil fuels as
was discussed under primary productivity.

a. To maintain early successional stages goes against the succession.

b. This is why we must plow and harrow the soil to create a nudation.

c. Plant in monoculture the edible plants and seeds for efficient harvests.

d. To protect them against pests, use pesticides and fungicides.

e. To eliminate the competing weeds, use herbicides.

IV. Paleosuccessions

A. Climatic change in the Cenozoic period

1. In the early Cenozoic about 60 million years ago, when the world was warmer
than now, an Arctotertiary forest covered Northern Eurasia and North America.
It was composed of a species-rich mixture of angiosperms and conifers of
similar overall species composition.

2. Southern North America was covered by a Neotropical Tertiary forest.

3. Through the Cenozoic, the past 60 million years, the climate has progressively
overall changed to a cooler drier climate and mountain building further
modified the climate drastically.

4. The Neotropical Tertiary forest was driven south to Central America. The
Arctotertiary forest was broken up and modified in different areas.
 5. A new flora, the Madrotertiary forest arose in the drier areas of the Mexican
Plateau and southwestern USA.

6. Further cooling led to the Pleistocene epoch in which there were cyclic
glaciations, at least four big glaciations, which drove the northern forest south
in both North America and Eurasia modifying the forest greatly.

7. Because the Mediterranean Ocean stood in the way and transverse mountains of
Europe spawned glaciers, the European sector suffered great extinctions of the
Arctotertiary Forest.

B. Plant fossils document these changes.

1. The most important fossils are pollen grains of the science of Palynology.

2. When they fell into dystrophic lakes and bogs, the acid conditions preserved
the pollen grains, which allowed a sequential record to be laid down, which
documented the changes in the vegetation over time.

3. Palynology reveals paleosuccessions of changes in plant communities, which

documents the drastic climatic changes that has occurred (see Fig 21.20 in
Smiths).

4. Shows that during the Pleistocene, the tropics were strongly affected in that
when the ice sheets were at there maximum, the equatorial area dehydrated and
the savannas spread at the expense of the forest (see Figs. 21.21 to 21.27 in
Smiths).

5. The wet tropical forests fragmented into refugia and it is reasonable to use this
fragmentation to explain the forces of geographical isolation that would
contribute to creating the high numbers of species in the tropical forests.

6. During interglacials (and some were warmer than the present one) the rainfall
increased in the tropics resulting in pluvial periods.

7. During times of high glaciations in contrast, the temperate deserts became

moist, a different kind of pluvial. Great Salt Lake is a remnant of the huge
Lake Bonneville that formed during the most recent glaciation, the Wisconsin.

8. Since the last glaciation, the climate has been progressively changing, and even
the last glaciation was only 12,000 years ago. It’s wrong then to think that any
of the ecosystems of the world have lasted for millions of years and
communities have been undergoing continual change for a long time, but this is
relatively slow change not the rapid changes that Mankind is bringing about
today.

C. Further back in time, in the Mesozoic and the Paleozoic, 230 and 600 millions of
years ago, palaeoecological extrapolations can be made (see Table 21.1).
1. Palaeoecology can be most successfully applied to marine environments, which
have fine records because these are primarily depositional environments,
favorable for preserving fossils especially of mollusks.

2. By comparing modern conditions and organisms to those of the past good

extrapolations can be made about palaeoclimates and past ecosystems.

3. Furthermore, geology can now age the sea bottom rocks, and measure the
magnetic polarity in rocks of land and sea, and so tell where a continent (or part
of a continent) was in space and time.