Lesson 20: Succession

• Definitions
– Seres and stages
– Autogenic vs. allogenic succession
– Progressive vs. retrogressive succession
– Facilitation, inhibition, life-history traits
• Examples of primary and secondary succession
• Methods of documenting succession
– Historical long-term data from permanent plots
– Examination of plots along a chronosequence
– Pollen record
– Dendrochronolgy
• Models of succession
• Facilitation vs. inhibition (Cowles vs. Clements)
• Relay floristics vs. intial floristic composition (Egler vs. Clements)
• Plant strategies vs. resource limitation (Grime vs. Tilman)
 Exam Friday, March 10
• The exam will cover material since that last exam, Lessons 12-21.
• I will send the exam out via email by 5 pm on March 10.
• The exam is due at 9 am on Tues Mar 21, following Spring Break.
• The format will be similar to the first exam except there may one or two short essays.
 Succession

• “The directional change in the species composition or structure of a plant community

over time.”

• It refers to the progressive change in the dominance and composition such that the
community will not return to former state without a disturbance that resets the
process of succession.

• Generally, it refers to the changes that occur with time scales of less than 500-1000
years (However, in the Arctic successional changes can be extremely slow, occurring
over periods of thousands of years, in the case glacial succession.)

• It does not include the regular seasonal changes in plant communities, nor the
regular changes that may be due to predictable climatic events (e.g., burst of annuals
following rain in the desert).

• It also does not include the extremely long-term change associated with geologic
changes (does include glacial succession), or long-term climate change.
 Cowles Chicago School of Dynamic Ecology
• Cowles and his students started the dynamic
approach to studying vegetation change through
time.

• Cowles (1899) study of succession on the sand

dunes around Lake Michigan was the first major
study of succession in North America.

• Cowles was a contemporary and admirer of Warming

Pond succession in Ohio: Dachnowski
who revolutionized botanical studies with his
famous work Plantesamfund in Danish in 1895. This
was later translated into German (Lehrbuch der
Ökologischen Pflanzengeographie) in 1896 and
finally into English (Oecology of Plants: An
Introduction to the Study of Plant Communities) in
1909. Warmings last chapter is devoted issues of
succession although he does not use this term.
 Seres and stages

• Sere: a series of plant communities that follow one another in an

ecological succession from a pioneer stage to, and terminates in a
particular stable (climax) association.
– Primary sere (prisere): A sere involving a primary bare surface that has not been
previously altered by vegetation, such as that occurring on rock, a recently
deglaciated surface, a new river bar, or pond.
– Secondary sere: A sere involving revegetation of a bare area resulting from
destruction of the previous vegetative cover, such as following a fire, cultivation,
or destruction by anthropogenic pollution such as smelter fumes.

• Stage: any floristically or structurally distinctive segment of a sere, (= a

seral stage).
Primary, secondary, and cyclic succession

Primary succession occurs on new newly

exposed land such as river floodplains, or
glaciated landscapes.

Cyclic succession can occur at any stage of a

sere due to periodic regular disturbance. But
the vegetation is not cleared.

Starting with new land, succession proceeds

through to the climax through a series of seral
stages.

A subclimax can occur when a periodic

disturbance, such as fire maintains a
vegetation at a point short of the regional
climatic climax.

From Daubenmire 1968

Secondary succession occurs when the
vegetation is cleared but the soil remains.
 Autogenic vs. allogenic succession

• Autogenic succession: The plants themselves cause succession to occur

(e.g. light capture by leaves, production of detritus, water and nutrient
uptake, nitrogen fixation).

– Facilitation: The process whereby site factors are improved through the process
of succession for establishment of the next sere (e.g., increased organic matter,
nitrogen fixation, moister soils for seedling establishment.)

– Inhibition: The process whereby factors inhibit the establishment of new

species, or growth of existing species (e.g., reduced light levels, competition
from other species). Factors can facilitative for some species and inhibit the
growth of others.

• Allogenic succession: Factors external to the plants cause succession

(e.g., climate change, periodic fires, floods).
 Examples of primary succession: Rock outcrop sere

Left -- Quartzite rock with Grimmia clones expanding from crevices. Crustose and Based on Oosting and Anderson 1939.
foliose lichens cover most of the surface elsewhere. In Daubenmire 1968.

Right - Successively wider rock crevices from right to left suggest stages of
succession from herbs to low shrubs to tall shrubs as the fracture planes weather. The
stabilized lichen-covered surface tends to be overriden by vascular communities
expanding from the crevices. In the background is a suggestion of a still later stage
consisting of conifer dominated forest. Cadillac Mountain, Mt. Desert Island, Maine.
Principal stages of primary succession on granite rock

Based on Oosting and Anderson 1939.

In Daubenmire 1968.
 Sand dune seres
Dune succession:
Dune succession has been studied by many ecologists in various parts of the world, including Warming in Holland and the famous study
of Cowles on the Michigan dunes.
The sand dune sere starts with a bare sand surface.
It is followed by an initial sand binder such as Elymus mollis or shrubs in the case of desert dunes such below.
The left photo are dunes captured by Abronia maritima and Frnseria in the Baja California.

Dunes captured by Abronia maritima and Fraseria in the Baja Dunes in the Salton Sea of California with Prosopsis
California. pubescens that has grown to keep pace with sand that collects
in the still air beneath the branches.

From Daubenmire 1968

 Dune stabilization

Dune surface near Minneapolis that has been quiescent for several Dunes well stabilized by Ericameria ericoides and other shrubs along
years, resulting in the establishment of many young plants of the the California coast near Santa Maria.
coarse Calamovilfa longifolia,

From Daubenmire 1968

 Hydrosere

Pond in northern British Columbia formed by

deglaciation several thousand years ago.
Competitive elimination and mutual adjustments
have produced narrow but sharply demarked
vegetation zones (Nymphaea, Carex,, Betula-
Salix, Picea mariana) on a gradual environmental
gradient.
Daubenmire, 1968
Sequence of dominant plant in a hydrosere of lower Michigan
(Gates 1942)
• The replacement of Carex
by Chamaedaphne marks
the transition from fen to
flat bog in which the ground
beneath the woody plants
has a carpet of Sphagnum
throughout. pH changes
from non-acidic to acidic.

Fen to bog
transition
 River floodplain sere: Tanana River
(Viereck 1970, Van Cleve and Viereck 1981, L. Walker et al., 1986; Adams 1999)

Diagram from Bonanza Creek LTER web page.

Diagram from Adams 1999.
Primary succession on Tanana River bars

Bonanza Creek Photos courtesy of Phyllis Adams.

Tanana River Succession, Stage I, bare surface
Tanana River Succession, Stage III, open shrub
Soil binders: rhizome system of Salix
Tanana River Succession, Stage IV, Closed shrub
Tanana River Succession, Stage V, young balsam poplar
Tanana River Succession, Stage VI, mature balsam poplar
Tanana River Succession, Stage VIII, mature white spruce
Tanana River Succession, Stage IX, black spruce
Progressive and retrogressive succession on Alaskan floodplains

From Drury, W.H. Jr. 1956. Bog

flats and physiographic processes
in the upper Kuskokwin River
region, Alaska. Contributions
from the gray Herbarium 178: 1-
30.
 Progressive vs. retrogressive succession

Progressive succession: Succession with increase in biomass,

diversity, and structural complexity through time, usually with a
trend toward a more mesic (moderate) site.

Retrogressive succession: Succession with a decrease in diversity

and biomass, often with the site becoming more hydric (wet) or
xeric (dry).
Some vegetation and ecosystem trends in progressive succession
Retrogressive succession
on old sea terraces in
California (Jenny 1969)

Involves development of
a soil hardpan and
water-logging of the
site (paludification).
Retrogressive succession involving bryophyte paludification in
southeast Alaska (Klinger 1990)
 Paludification

The process of mire (peat-forming ecosystem) formation over previously

forested land or grassland due to climatic or autogenic processes leading
to waterlogging and anaeroby.
Effects of bryophyte interactions on soils and vegetation
during succession (Klinger 1990)
 Secondary succession following grassland fire

Edge of a burned area which was

burned the preceding August and
photographed in May. In this stable
community dominated by Bromus
tectorum, small changes in the
relative development of species
are conspicuous the first postfire
season, but negligible thereafter.
Dry matter production increased
temporarily after burning.
Lewiston, Idaho. Npte the two
researchers working along a
vegeation transect.

Photo from Daubenmire 1977

 Succession in grasslands following fire

• Frequent fire is the agent for soil-nutrient renewal. Many nutrients

are locked in the abundant litter until fire occurs.
• C4 (warm-season) grasses are at advantage in fire dominated
systems, and will out-compete C3 grasses and forbs.
• Mechnaisms for survival:
– Dominance of hemicryptophytes with perrenating buds at or below the
ground surface (lethal temperature for most plants is 60˚C). The plants
die back annually creating abundant standing dead vegetation and
litter, which is an ideal fire fuel.
– Increased reproduction from seeds. Many seeds require scarification
(any agent that ruptures the integrity of the seed coat, leaving it
premeable to water and oxygen).
Secondary succession following forest fire

Daubenmire 1977
 Forest adaptations to fire

• Serotiny: Late-to-open cones that retain viable seeds (sometimes

for decades) until some mechanism severs the vascular
connection with the parent plant, or breaks the resinous bonds
that seal the cone scales.
– Seedling recruitment is disturbance dependent.
– Seeds are not resistent to heat so there is little or no persistent soil
seed bank.
– Seeds are shielded from excessive heat by woody fruits or cones.
– Late summer or early fall fires are best for seedling survival.
• Thick fibrous bark.
• Evanescent lower branches.
• Rapidly growing seedlings.
 A general fire hypothesis

“For those ecosystems in which fire occurs predictably and frequently

enough to result in a degree of fire-dependence, fire serves to increase
and/or maintain the availability of essential resources which would without
fire be growth-limiting for the organisms in the systems”
Gilliam 1991,
The significance of fire in an oligotrophic system
 Some general properties of fire-maintained systems

• Plants have properties that make them susceptible to burning including

the arrangement of plant parts in space, as well as the size and shape of
those parts.
– Fine, xeromorphic leaves and twigs, with high surface-to-volume ratios.
– Quick to dry out.
– Favorable fuel-to-air ratio.
– Accumulation of fuel with low moisture (<5-15% moisture in dry weather).

• In soil seed storage, with fire stimulated germination.

• Fire induced flowering.

• Bud protection and sprouting subsequent to fire.

• On-the-plant seed storage and fire-stimulated seed dispersal.

• Trees with thick bark.

• Plants with rapid growth and early maturity.

 Secondary succession: Old agricultural fields in North Carolina
(Based on Oosting 1942)

Conyza canadense and Digitaria sanguinalis dominating an abandoned field on which

Zea mays was grown the preceding year. Florence South Carolina.
Photo from ; Photo from Daubenmire 1977 1977
Old field succession: 3-6 yr after abandonment

Old fields
dominated by
broomsedge
(Andropogon
virginicus)
with scatterd
pine seedlings
are typical of
secondary
succession 3-
6 yr after
abandonment
and prior to
dominance by
pine.

From Colinveaux 1973, in Barbour et al. 1999, p. 286.

 Old field succession: 50 yr after abandonment

A loblolly pine stand with hardwoods understory, typical of secondary

succession in the Piedmont.
Secondary successional trends in dry and moist abandoned fields,
North Carolina (Based on Oosting 1942)
 Secondary succession: Density of pine and hardwood overstory
and understory species after 100 yr in North Carolina (Oosting 1942)

• Decline of pines and increase in hardwoods toward climatic

climax.
Cyclic succession in a Texas desert (Yeaton 1978)

• Opuntia leptocualis
uses Larrea tredentata
as a mother plant.

• Eventually Opuntia
outcompetes Larrea.

• Until Opuntia dies

creating an open patch
for Larrea to colonize.
 Methods of documenting succession

• Historical long-term data from permanent plots

• Examination of plots along a chronosequence
• Pond sediment, pollen records
• Dendrochronolgy
Methods of examining succession: Serial mapping
Example: 3 yr period in eastern Nebraska during a severe drought.

From Weaver and Alberton (1936)

Repeated measures documenting succession in aspen forests,
Michigan (Roberts and Richardson 1985)

Stand
ages:
a. 20
b. 30
c. 35
d. 49
e. 60

• Trees follow self thinning law during succession.

• Number of all stems declined until 55 years.
• Basal area increased most dramatically in aspen and red pine.
 Evidence of succession in pond sediments (Rigg and
Richardson 1938)

Stratification of peat types in

a bog near Puget Sound.
The stratigraphy shows a
regular and uninterrupted
sequence of stages such
as might be inferred from
the present community
zonation about ponds in
that region.

From Daubenmire 1977, p. 144.

 Pollen Diagrams

• Pond succession, Waupaca

bog, Wisc.
• 23 feet of peat.
Interpretation:
• Picea-Abies forest
surrounded the pond
during early succession.
• Pinus and Quercus
replaced the spruce-fir as
the pond filled with limnic
sediments.
• Temporary rise of oak at
12 ft indicates warm
climate.
• Recent cooling saw the
advance of pine over oak.
• A sedge peat developed
after the lake filled with
sediment.
• Sphagnum invaded as the
Voss 1934 pH of the site declined.
Population structure
 Size classes of trees:
Which trees are coming and which are going?
 Succession theory: Clements vs. Cowles

• Clements saw succession as a predictable process with definite

“seres” or stages of succession. In one sense he saw the
community as a “superorganism”, with predictable stages of
development. In other words, autogenic processes predominate.

• Cowles saw succession as much more dynamic, a process that

could be influenced by innumerable outside forces that could lead
to different end points depending on, for example, circumstances
of propagule availability, weather and climate during succession,
and factors of the microsite. In other words, allogenic processes
predominate.
 Models of succession: “Relay floristics” or “facilitation”
This is a diagrammatic
summary of relay floristics,
Frank Eglers view of
Clementsian succession.
He viewed this as a
concept where one group
of species was replaced by
another group, each
representing a distinct
“stage” of a sere.

This view has also been

termed facilitation by
Connell and Slatyer (1977).

In this view, one stage of the

sere prepares the habitat
for introduction by the next
stage, often through
changes in soil (e.g.
nutrient additions) or by
otherwise making the site
more conducive to the next Based on Egler 1954
group of species.
“Initial floristic composition” model of succession
(Egler 1954)
• Egler challenged the facilitation
or relay floristics model by
stating that the patterns of
succession are more often a
function of life-history patterns.
• For example, in old field
succession, most of the
pioneering species, many of the
seral species and a few of the
climax species are present
before abandonment of the field.
• Some germinate quickly and
express themselves early in the
successional. Others express
themselves later.
• Longer living, larger, slower
growing plants eventually
outcompete smaller pioneer
species.
• Community dominance shifts
and succession proceeds.
 Grime’s triangular model of succession • Plant strategy diagram
predicts how plants with
various strategies (R, C,
S) would respond to
Potential productivity succession.
• Diagram shows how the
P1 = High (e.g., fertile soils) abundance of various
P2 = Moderate plant strategies would
P3= Low (e.g., infertile soils) change under
succession in differing
conditions of soil
Size of circles represent fertility.
amount of biomass. • In all situations,
Ruderals (R-strategy
plants) initiate the
process of succession,
The abundance of C-
and S-strategy plants
increase through
succession.
• P1 shows the situation
with high soil fertility.
• P3 is the situation under
low fertility when
biomass potential is
low, and stress
tolerators are the main
survivors.
• C -strategy plants
(competitors) play a
major role particularly in
the middle portion of
each sere.
Trajectories of succession using Tilmans resource ratio hypothesis

• Diagram show how

various combinations of
species might occur with
different starting points of
nutrients in the soils.

• This model predicts rapid

succession (few seral
stages shown by the
numbers) with minimal
species change on fertile,
productive soils, and slow
succession with greater
species change on
infertile, less productive
soils.
 Primary succession: Glacier Bay, Alaska
(Cooper 1923-39; Crocker and Major 1955; Lawrence 1967; Chapin et al. 1994)

One of the classic sites for study of

succession has been the study of the
vegetation changes following the
retreat of the glaciers in Glacier Bay,
Alaska. John Muir and many others
commented on the the obvious
patterns of succession in the Bay.

William Cooper, one of Cowles

students, began formally studying this
area in 1920s, and numerous others
have followed, including Terry Chapin.
 “Mechanisms of primary succession following deglaciation at
Glacier Bay, Alaska” (Chapin et al. 1994)

Examined the relative importance of facilitation, life-history traits, and

competitive interactions at a site that has been used as the classic
example of facilitation (Cooper 1939; Crocker and Major 1955; Lawrence
1951, 1959, etc.).
Map of Chapins Glacier Bay study areas
 Pioneer stage
From Chapin et al. 1994

Aerial view of pioneer stage (a) at the time of

deglaciation in 1974. Arrows indicate borders of
the the study area about 2 km apart

Beginning of the study in 1987. A shade

frame for the seedling experiment and a plot
from which the black crust has been removed
for the seed-sowing experiment are shown in
the foreground.
Beneath the McBride Remnant in 1949, (center of
photograph).

Dryas
stage

Following deglaciation in 1967

At the beginning of the study in 1986

Chapin et al. 1994. Photos

from 1949 and 1967 courtesy
of D.B. Lawrence
 Alder stage

(a) At the time of deglaciation

in 1929 (arrows indicate
borders of the study area.

1929 1949 (b) During transition from

pioneer to the Dryas stage in
1941.
(c) at the beginning of the study
in 1986.
(d) View of the study area on
the far (south) side of
Nunatak cove in 1949,
(e) 1967
(f) 1987.

1941 1967

Chapin et al. 1994. Photos 4b, d,

and e courtesy of D.B.
Lawrence.

1986 1986
Spruce stage
 Study Methods (Chapin et al. 1994)
• Site characterization at 10 study plots within each successional stage
(soil, nutrient pools, PAR) (facilitation study).
• Greenhouse plot tests to ascertain changes in soil fertility (facilitation
study).
• Life history traits of Dryas, alder, and spruce within each successional
stage (seed weights, growth rates, age of first reproduction, maximum
longevity, maximum height) (life history study).
• Seed arrival and germination (seed traps, in each stage, seedling
transplants of alder and spruce in each stage, studies of seedling
biomass) (life history study).
• Examination of natural seedlings growth (life history study).
• Experimental seedling transplants of alder and spruce (shade
experiments, litter experiments (inhibition experiment).
• Alder litter experiment on Dryas growth (inhibition experiment).
• Allochemical effect of plant litter on the establishment of seedlings and on
growth of the key species (inhibition experiment).
 Soil properties during primary succession, Glacier Bay, AK (Chapin et al.
1994)

• Invasion by trees in the Spruce

Stage triggers increased soil
moisture, increased N, and
decreased bulk density, pH, and
P.
• Changes to the moss layer
caused most of these changes.
 Biomass of spruce seedlings during succession, Glacier Bay, AK (Chapin et
al. 1994)

• Spruce seedlings were most

abundant and had greatest
biomass during the Dryas
stage.
• At later stages the seedlings
were inhibited mainly due to
shading effects.
Facilitative vs. inhibitory effects of each stage on establishment of
spruce seedlings
Conclusions: Succession is a function of a combination of facilitation,
inhibition, life-history traits, and stochastic factors
(L. Walker and Chapin 1987, L. Walker et al. 1994)

*Life history traits: Such factors as size, rate of growth, seed size, and
dispersal are of primary importance and control the general patterns of
succession. Proximity to seed sources are very important.

Facilitation: The rate of these patterns is greatly enhanced by changes to the

site such as addition of nutrients, organic matter, and soil moisture which
facilitate the growth of species in later stages of succession.

Inhibition: Also affects the rate through competitive interaction between

species or through allochemicals that may inhibit growth of other species.

Stochastic factors affecting the site: Unpredictable abiotic events such as

climate change, floods or catastrophic events can reset or change the
direction of succession.
 Lesson 20: Summary
• Definitions
– Seres and stages
– Autogenic vs. allogenic succession (Clements vs. Cowles)
– Progressive vs. retrogressive succession (Drury 1956; Klinger 1990)
– Facilitation, inhibition, life-history traits (e.g., Chapin et al. 1994)
– Cyclic succession (e.g., Yeaton 1978)

• Examples of primary and secondary succession (Tanana River, Viereck

1970, Van Cleve and Viereck 1981, Walker et al., 1986; Adams 1999; Glacier
Bay (Chapin et al. 1994); North Carolina old fields (Oosting 1942); fire
succession

• Methods of documenting succession

– Historical long-term data from permanent plots
– Examination of plots along a chronosequence
– Pollen record
– Dendrochronolgy

• Models of succession
– Facilitation vs. inhibition (Cowles vs. Clements)
– Relay floristics vs. intial floristic composition (Egler vs. Clements)
– Plant strategies vs. resource limitation (Grime vs. Tilman)

• Glacier study of Chapin et al. (1994) examined the relative roles of life
history traits, facilition and inhibition.