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Succession (遷移)

Mount Usu / Sarobetsu post-mined peatland
From left: Crater basin in 1986 and 2006. Cottongrass / Daylily

Changes in communities with time

Pioneer (先駆種): earlier successional species

tending to be sun tree and/or generalist

Climax species (極相種)

tending to be shade tree and/or specialist


Sussceesion is translated into 'sen-i (センイ, 遷移)'
[ secondary succession | Usu | chronosequence | facilitation ]

The mechanisms of succession explained by Grime's triangle (グライムの三角形による遷移機構の説明)
Fig. successional trajectories explained by Grime's triangle. Initial condition of productivity is important for those trajectories.

Theories of succession (Finegan 1984)

Emergent properties
Autogenic changes - facilitate succession
Orderly, predictable, deterministic
Individuals, or species
Autogenic changes inhibit succession
The sequential physiognomic dominance of
the site by species with different life histories,
growth rates, and size at maturity
Clements (1916)

Monoclimax theory - climatic climax
Action, reaction, coaction

Tansley (1935): Polyclimax theory

Autogenic succession
Allogenic succession

Odum (1971)
Gleason (1926): Individualistic concept of vegetation

Whittaker (1953): Climax pattern theory
Egler (1954): Relay floristic composition
Drury and Nisbet (1973)

Initial floristic (mainly old field)

Connel and Slatyer (1977)

Rejected facilitation (reaction)
Accepted tolerance (mainly old field)
Accepted inhibition (mainly old field)

Monoclimax theory
Climatic climax (気候極相)
Polyclimax theory
Edaphic climax (土壌極相), determined by (poor) soil
Physiographic climax (地形極相), by topography
Biotic climax (生物極相), by life activities, such as grazing
diversity change
Changes in species diversity across succession (Drury & Nisbet 1973)


Primary succession (一次遷移)

Xeric succession (乾性遷移)
Case: volcanic succession (火山遷移), e.g., on Mount Usu and Mount St. Helens (セントへレンズ山) Schema of succession
Representative of primary, xeric succession (Whittaker 1975). Vegetation becomes tall with increasing time.
Mesic succession (湿性遷移)
Fig. Inundated or saturated so as to support or preduce of vegetation typically adapted to like in saturated soil.
Secondary succession (二次遷移)
Secondary succession: started by disturbances that do not completely remove the former preexisting vegetation and soil

Post-mined peatland
Revegetation on skislope

landscape of skislope
problems on skislope establishment
skislope establishment

Revegetation after coal mining
Revegetation after wildfire

Secondary succession (二次遷移)

In the early stages
2nd1 2nd2
[1/2] The landscape is one year after a bare land was created due to closing a pachinko parlors. The dominant species were Chenopodium album, Oenothera biennis and other annuals. The vegetation recovery is greatly determined by seedbank. On N24/E1, Sapporo City, on August 30 2008.
2nd3 2nd4
[3/4] These are the landscape of two years after the creation of bare land. Species composition changed greatly to Lactuca scariola (annual to biennial). [3] A perennial Artemisia montana patch developed. [4] Brown stalks are Rumex obtusifolius (perennial). On July 30 2009.
Secondary vegetation (二次植生): vegetation that develops after disturbances (s.l.)

secondary vegetation made by human activities (代償植生) (s.s.), e.g., agricultural lands, artificial forests and satoyama

Secondary forest or second-growth forest (二次林): forests out of secondary vegetation types

↔ old-growth, primary or primeval forest

Climax (極相)

Three factors on climax
  1. Stability: self-maintaining = mature plant community
  2. Convergence: final stages of succession in a given area
  3. Regional prevalence: characterized as principal, undisturbed plant community in a given region
    directional change → succession
    non-directional change → cyclic change or fluctuation

Successional pace (遷移速度)

Type I. Successional pace calculated along a temporal flow
1) Sørensen's community coefficient (遷移群落類似度指数), CC (Bornkamm 1981, Tsuyuzaki 1991)

CC = 2a/(2a + b + c)
a: number of species observed in both the times (years), b: number of species observed only in the first time, c: number of species observed in the next time

2) Percent similarity (遷移百分率類似度), PS (Bornkamm 1981)

PS = 2Σmin(xi, yi)/Σ(xi + yi) × 100
xi, yi: percentage cover of ith species in times (years) x and y, min(xi, yi): minimum percentage cover of ith species in the two years

→ basically these indexes are derived from similarity indexes
3) Speed of succession, D (遷移速度)

D = Σi=1s[n(x, y, t2) – n(x, y, t1)]2
s: number of species present within the plots,
ni: within-quadrat average covers of the species
x, y: grid coordinates

        Species  t1   t2  t2 - t1  Square
        1        10  20     10      100
        2         5   5      0        0
        3         .   5      5       25
        4         1   .      1        1
        5         1   .     -1        1
        Sum                         127

D = 127

4) Rate of succession (遷移率) (Prach et al. 1993)

Vi = maxj=1n(|PijP(i–1)j|)
for i = 2, …, N and species j = 1, …, n. The n is number of species considered in the respective sere (being dominant at some time) and N is number of years. Values Pij refer to the cover of a species in the first year and Pij generally to the cover of a species j in the i-th year.

Type II. Succession degree calculated by vegetation structure
1) Degree of succession (遷移度), DS (Numata 1969)

DS = (Σi=1nlidi)/n × v
l: longevity (annual or therophyte = 1 (biennial = 2), perennial (geophytes, hemicryptophytes, chamaephytes) = 10, nanophanerophytes = 50, megaphanerophytes = 100)
d: relative dominance: (C' + H')/2 or (F' + D' + C')/3
n: number of species in the minimum area
v: cover percentage, 100% = 1
Empirically DS is effective between the successional stages of grasslands and sun tree forests. However, DS does not indicate succesional stages well at the late successional stages becasue DS does not change greatly due to the dominance of N, M and MM.

2) Successional index (遷移指数), SI (Nakamura 1984, 1985)

SI = cΣi=1n(di·l)
d: relative dominance of basal area (range: 0-1), l: estimated tree age in the dominat stand on each species, c: crown area (100% = 1) d : relative dominance based on basal area (ranging from 0 to 1); l: life span which was estimated by the ages of massive stems of each species or species group in its dominating stands; c: cover of canopy (ranging from 0 to 1, where means 100% in cover) which would affect shade-tolerances of undergrowth).

Chronosequence (クロノシークエンス)

The sequential set of changes in structure and composition of (plant) communities.
In Japanese, there has not been any nice translation on chronosequence. The phonological translation is often used. Here, I indicate a tentative translation.


Plant communities on post-fire forest, glacier retreat, lava, skislope
All ages on the researched sites should be determined.

Chronosequence Fig. Hypothetical procedures to obtain successional sere by chronosequence approach. This is the case of post-fire stand. [Above] Circles indicate the areas that received fire. Squares indicate plots established in the post-fire areas. Numerals indicate the ages after fire. [Below] The sequential changes in species composition. Species A is dominant five years after fire, and then gradually decreases the abundance. Species B is the most common in area where it has passed for 20 years since the last fire. If we assume that the species composition change in time, we determine species A is a pioneer after fire in the region.

Ex. Sakurajima Island, Kyushu, Japan (Tagawa 1964)
Ex. skislopes (Tsuyuzaki 2005)

Problems on the assumption of chronosequence

  • The intensity, scale and frequency of disturbances are not always same.

[facilitation on Mount Koma]

Facilitation (定着促進効果)

Facilitation vs competition

Stress gradient hypothesis, SGH (ストレス勾配仮説)

Fig. Stress gradient hypothesis (Bertness & Callaway 1994)
often estimated by relative neighbor effect

(Maestre et al. 2009)

Revised SGH
Life history of interacting species

stress tolerant

Stress gradient type

resource (e.g., nutrients)
non-resource (e.g., frost)
Fig. Revised SGH

(Successional) facilitation

In which the influence of early species in a community succession is to facilitate establishment of later ones by changing the conditions encountered. (Begon et al. 1996)
Fig. Temporal changes in facilitation, competition and tolerance along successional sere

Types of facilitation


A. Ameliorate harsh environmental conditions = Resource modification

  1. Adjust light and temperature by shading
  2. Increase soil moisture
  3. Increase soil nutrients
  4. Adjust soil aerobic conditions
patch Fig. 2 Study site to evaluate the effects of two patch-forming shrub species, Salix reinii and Gaultheria miqueliana, on cohabitants on Mount Koma (photo by S. Uesaka). (Uesaka & Tsuyuzaki 2004). A deciduous shrub, S. reinii is likely to be facilitative, and an evergreen shrub, G. miqueliana seems to be inhibitive. The patches increase moisture and nutrients in the volcanic deposits, and decrease solar radiation on the ground surface. Tussocks (谷地坊主) in wetlands often have facilitative effects (Koyama & Tsuyuzaki 2010).

B. Alter the characteristics of soil substrates

  1. Bulk density, and macropores
  2. Nurse logs
  3. Seed trap effect by diverse microtopography

C. Epiphytes (e.g., orchids, ferns, mosses, algae)

  1. Autotrophic: obligate, facultative
  2. Heterotrophic: parasitic (asymmetrical effect)
  3. Auto-heterotrophic continuum (hemiparasites)
  1. Eliminate potential competitors
  2. Introduce beneficial organisms, e.g., mycorrhizae and pollinators
  3. Protect from herbivores by physical modification of environment


  • Tsuyuzaki, S. 1987. Origin of plants recovering on the volcano Usu, northern Japan, since the eruptions of 1977 and 1978. Vegetatio 73: 53-58
  • Tsuyuzaki, S. 1989. Analysis of revegetation dynamics on the volcano Usu, northern Japan, deforested by 1977-78 eruptions. American Journal of Botany 76: 1468-1477
  • Tsuyuzaki, S. 1994. Fate of plants from buried seeds on volcano Usu, Japan, after the 1977-78 eruptions. American Journal of Botany 81: 395-399
  • Tsuyuzaki, S. 2009. Causes of plant community divergence in the early stages of volcanic succession. Journal of Vegetation Science 20: 959-969
  1. Give some examples of arrested succession and discuss the mechanisms.
  2. A city lot is undergoing succession. Your goal is to reestablish a forest community. Under each of the following (example) conditions, what would you do? a) succession has been arrested by strong dominance of an exotic shrub such as Scot's bloom; b) there are serious barriers to immigration; c) soils have become compacted over the years; d) all of the above. Assess the nature of the succession if left alone, then determine if you will be accelerating, deflecting, or correcting the succession tract. Give brief examples of your tactics.
  3. Name some stochastic events and give examples about how they might alter succession.
  4. Contrast facilitation with inhibition as succession mechanisms. What does each predict concerning the order and structure of succession? Are these concepts mutually exclusive?
  5. List five major factors or processes that would produce a plant association distinct from the climatic climax type in an region where you like.
  6. List five factors that have influenced the landscape of Lake Toya.