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2. Community Ecology and Dynamics –
Succession and Stability
H.J.B. Birks
BIO-201 ECOLOGY
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Community Ecology and Dynamics - Succession and Stability
Some ecological and environmental basicsSuccession Basic concepts
Primary succession on glacial forelandsCommunity changesEcosystem changesMechanisms of succession
Stability Basic conceptsWhat causes resilience?Alternative stable states and regime shiftsMaintenance dynamicsDisturbance and diversity
Community concepts revisitedConclusions and Summary
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Pensum
The lecture, of course,
and
the PowerPoint handouts of this lecture on the BIO-201 Student Portal
Also ‘Topics to Think About’ on the Student Portal filed under projects
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Topics to Think About
On the Bio-201 Student Portal filed under Projects, there are several topics to think about for each lecture. These topics are designed to help you check that you have understood the lecture and to identify important topics for discussion in the Bio-201 colloquia.
In addition, there are two or three more demanding questions at the sort of level you can expect in the examination question based on my 10 lectures. These can also be discussed in the colloquia.
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Background Information
There is now a wealth of good or very good ecology textbooks but perhaps no excellent, complete, or perfect textbook of ecology.
Not surprising, given just how diverse a subject ecology is in space and time and all their scales.
This lecture draws on primary research sources, my own knowledge, experience, observations, and studies, and several textbooks.
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Textbooks that provide useful background material for this lecture
Begon, M. et al. (2006) Ecology. Blackwell (Chapter 16, 1 in part)
Bush, M. (2003) Ecology of a Changing Planet. Prentice Hall (Chapters 15, 16)
Krebs, C.J. (2001) Ecology. Benjamin Cummings (Chapter 21)
Miller, G.T. (2004) Living in the Environment. Thomson (Chapter 8)
Molles, M.C. (2007) Ecology Concepts and Applications. McGraw-Hill (Chapter 20)
Ricklefs, R.E. & Miller, G.L. (2000) Ecology. W.H. Freeman (Chapter 28)
Smith, R.L. & Smith, T.M. (2007) Ecology and Field Biology. Benjamin Cummings (Chapters 21, 22)
Townsend, C.R. et al. (2008) Essentials of Ecology. Blackwell (Chapters 9, 10)
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A Reminder
If you try to read Begon, Townsend, and Harper (2006) Ecology – From Individuals to Ecosystems, there is a 17-page glossary of the very large (too large!) number of technical words used in the book on the Bio-201 Student Portal. It can be downloaded from the File Storage folder.
Good luck!
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Some Ecological and Environmental Basics
Environment varies continuously in SPACE at all spatial scales (geology, soils, climate, altitude, slope, etc.) and varies at all TIME scales (days, months, seasons, years, decades, centuries, millennia, etc.)
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Broad spatial scale
15,000 ft10,000 ft5,000 ft
Coastalmountainranges
SierraNevadaMountain
GreatAmericanDesert
RockyMountains
GreatPlains
MississippiRiver Valley
AppalachianMountains
Coastal chaparraland scrub
DesertConiferousforest
Coniferousforest
Prairiegrassland
Deciduousforest
Average annual precipitation
100-125 cm (40-50 in.)
75-100 cm (30-40 in.)
50-75 cm (20-30 in.)
25-50 cm (10-20 in.)
below 25 cm (0-10 in.)
Biomes
Role of climate
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Long time scales
a) Change in temperature in the North Sea over the past 65 million years (M yr).
b)The ancient continent of Gondwanaland began to break up about 150 M yr ago.
c) ~50 M yr ago distinctive bands of vegetation had developed.
d)By 32 M yr these are more sharply defined.
e) By 10 M yr ago much of the present geography of the continents was established but with different climates and vegetation from today: position of Antarctic ice cap is schematic.
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Changing continental positions in last 220 million years
Tectonic plates in constant motion. Environment on earth changes accordingly.
1. Triassic220 million years ago
Pangaea continent had its maximum size. Large interior areas, very dry and extensive deserts.
2. Mid-Late Jurassic155 million years ago
Beginnings of the break-up of Pangaea.
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3. Late Jurassic149 million years ago
Break-up of Pangaea, large (100 m) rise in sea-level, Siberia and China now island continents, Europe a series of islands.
4. Early Cretaceous127 million years ago
Break-up of Gondwana.
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5. Mid Cretaceous106 million years ago
Europe still a series of islands, North and South America widely separated.
6. Late Cretaceous65 million years ago
Similar to today but for North and South America and India.
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CryogenianCryogenianCryogenian/Neoproterozoic IIINeoprotoerozoic IIINeoproterozoic IIICambrianOrdovicianSilurianDevonianCarboniferousPermianTriassicLate TriassicJurassicCretaceousPalaeogeneQuaternary
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At the same time, major changes in plant evolution and hence in earth vegetation
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Major evolutionary developments in last 500 million years
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Global ecological changes in the last 55 million years
1. Eocene55 million years ago
Widespread tropical rain-forest and no ice-caps
2. Late Eocene35 million years ago
Cooler, less tropical rain-forest, some ice-caps
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3. Oligocene25 million years ago
Cooler, more extensive Antarctic ice-cap. Semi-arid scrub and desert areas, evolution of giant land mammals
4. Miocene3.2 million years ago
Continents almost in today's position, ice-caps at both poles, climate drier, vast grasslands, much mountain uplift
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5. Late Pliocene1-2 million years ago
Extensive polar ice-caps, much reduced tropical rain-forest
6. Pleistocene30 000 years ago
Massive ice-sheets, much tundra and arid vegetation
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Temperature changes in the Northern Hemisphere at different time scales
Shorter time scales
102
103
104
105
5x105
years years
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End of LIA
Medieval optimum LIA
Holocene
11500 years
Last millennium
LIA = Little Ice Age
Past 130 years
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Medieval Warm Period
LIA
Millennium scale: warm period 1000 AD and the Little Ice Age
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BiosphereBiosphere
Ecosystems
Communities
Populations
Organisms
Biosphere
Biomes
Ecosystems & Landscapes
COMMUNITIES
Species
Populations
Organisms
Today’s Ecological Scale
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Succession – Basic Concepts
1.Changing plant and animal communities, ecosystems, and landscapes through time following the creation of new substrates or following disturbance, usually directional changes.
2.Primary succession – occurs on newly formed surfaces such as volcanic lava flows, areas recently deglaciated (glacial forelands), sand-dunes along coast, etc.
3.Secondary succession – occurs where disturbance destroys a community without destroying the soil. Occurs after agricultural areas are abandoned, after forest fires, forest clearance, erosion, etc.
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4. Successional change is usually directed towards the undisturbed surrounding vegetation and fauna.
5. Succession generally ends with a mature community whose populations are relatively stable. 'Climax vegetation'.
6.Environment is changing at a range of scales in time and space, so communities are always in a state of flux and change.
7. Successional time scales – can be short or long. Few years; 250 years after the Little Ice Age; 10000 – 11500 years since the last glaciation.
8. Ecological succession “non-seasonal, directional, and continuous pattern of colonisation and extinction on a site by species populations” (Begon et al. 2006 p.479)
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Primary succession
Time
Small herbsand shrubs
Heath mat
Jack pine,black spruce,
and aspen
Balsam fir,paper birch, and
white spruce
Exposedrocks
Lichensand mosses
e.g. New surfaces formed by:
Glacier retreatVolcanic eruptionCoastal sand-dunes
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Secondary succession
Time
Annualweeds
Perennialweeds and
grasses
ShrubsYoung pine forest
Mature oak-hickory forest
e.g. Disturbance by:FireForest cuttingErosionWind-throw & storms Abandoned fieldsLarge herbivores e.g. elephants
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Differences between primary and secondary succession
Primary succession: no soil, no seed-bank, no organic matter
Secondary succession: soil is present but disturbed, seed bank present, organic matter present
Secondary succession is very common within landscapes, primary succession is less common
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Primary Succession and Glacial Forelands
Little Ice Age at about 1750 AD caused rapid advance of glaciers in, for example, Jostedal and Jotunheimen.
As ice subsequently retreated, deposited glacial moraines (silt, sand, gravel) on which primary succession could begin.
Some classic studies mentioned in this lecture:Nigardsbreen, Jostedalsbreen - Knut FægriStorbreen, Jotunheimen - John MatthewsKlutlan Glacier, Yukon - John BirksGlacier Bay, Alaska - W. Cooper et al.
Surface ages determined by historical observations, from the size of lichen (lav) thalli on rocks on the surface ('lichenometry'), and from annual growth rings of shrubs and trees. Surfaces of different ages form a CHRONOSEQUENCE.
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Chronosequences – series of sites (e.g. glacier moraine forelands, volcanic lava flows, sand dunes, recently formed islands) of different but known age.
Study vegetation and soils today on surfaces of different but known age.
Substitute space today for time – "space-for-time" substitution.
Glacier Moraines
Age of formation
1930
1890
1850
1750
soil pH
distance from glacier Age
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Nigardsbreen 'Little Ice Age' moraine chronology
Photo: Bjørn Wold
Knut Fægri
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1750
1770
1815
1912-30
Mature Alnus forest
Mature Betula forestMature
Betula forest
Photo: 1984
Primary Succession after Little Ice Age
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Nigardsbreen, Jostedalsbreen
1874 1931
1900 1987
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Nigardsbreen, Jostedalsbreen
2002
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Vegetation changes since ice retreat
20 years 80 years
150 years 220 years
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Styggedalsbreen, Jotunheimen
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Distribution of selected species on Storbreen moraines
‘Pioneer’ r-selected species
‘Late stage’ K-selected species
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Klutlan Glacier, Yukon
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Moraines of different ages at the terminus of the Klutlan Glacier
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Pioneer plants on Moraine II (2-5 yr) (Crepis nana)
Dryas drummondii mats (9-25 yr)
Moraine II (10-30 yr) Moraine III (30-60 yr)
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Moraine IV (60-80 yr)
Moraine IV (95-180 yr) Moraine V (180-240 yr) Harris Creek (>250 yr)
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Species abundance change with time
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Changes in major plant-growth forms with time
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Glacier Bay, Alaska
Phases
1. Pioneer phase – 20 years – Epilobium latifolium, Dryas drummondii, Salix spp.
2. 30 years - Dryas mats with Alnus crispa, Salix, Populus, and Picea
3. 40 years – Alnus forms dense thickets
4. 50-70 years – Picea and Populus grow above Alnus
5. 75-100 years – Picea forest with mosses
6. 200 years – Tsuga heterophylla & T. mertensiana forest
7. >300 years – more open forest with areas of bogs and tundra meadows
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Some Glacier Bay pioneer species
Dryas drummondii
Epilobium latifolium William S.
Cooper
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1957
2002
Little Ice Age maximum
Little Ice Age in Nepal about 1850
O.R. Vetaas
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river
Glacial lake
Lateral moraine stages
Terminal moraine-complex
Little Ice Age maximum (app. 1850)
Neoglacial stages (> 1200 BP)
1988
2001
Glacier fronts
Glacier in 1957
Gangapurna North Nepal stages since 1850 to present
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Lateral moraines with trees, Gangapurna, Nepal
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Other Primary Successions
1. Coastal fore-dunes
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2. Volcanic lava flows
Craters of the Moon, Idaho
Plant colonisation
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Over time species invade, then increase, some decrease again and disappear, and some remain as the mature vegetation
pioneerspioneers & late-invaders
Late invaders
Woody & long lived species
TIME
1. Changes in plant abundance and species composition in primary succession
Community Changes During Succession
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Late-succession species
K-selected
Fewer, larger offspring
Short dispersed seeds
Later reproductive age
Most offspring survive to reproductive age
Lower population growth rate (r)
Adapted to higher nitrogen and low light (shade)
High ability to compete
Early-succession species
r-selected
Many small offspring
Far dispersed seeds
Early reproductive age
Most offspring die before reaching reproductive age
High population growth rate (r)
Adapted to low nitrogen and high light
Low ability to compete
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2. Changes in species richness in primary succession over 1500 years
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Over longer time scales (> 2000 yr) richness often declines. Why?
Successional time
Speci
es
rich
ness
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Succession of plant growth forms at Glacier Bay
3. Changes in plant growth forms in primary succession
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Time
Annualweeds
Perennialweeds and
grasses
ShrubsYoung pine forest
Mature oak-hickory forest
4. Changes in species richness in secondary succession from 80 days to 200 years
Eastern N America – abandoned fields, tree colonisation and forest development 200 yearsSoil and buried seed bank present at the outset
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Number of breeding bird species
Woody plant species richness
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Number of macroinvertebrate and macroalgae species during secondary succession
Rocky coastal shores: 18 months
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Algal species diversity during secondary succession
Rivers after extreme floods: 80 days
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5. Species replacement during secondary succession
Henry Horn – predictive model for changes in tree composition given
(1) for each tree species, probability that within a particular time, an individual would be replaced by another of the same species or by a different species
(2) an assumed initial species composition
Horn argued that the proportional representation of various series of saplings established beneath an adult tree reflects the probability of that tree’s replacement by the species represented by the saplings.
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Using this, Horn estimated probability after 50 years of a site occupied by a given species will be replaced by another species or will still be occupied by same species in a forest in New Jersey, USA
A 50-year tree-by-tree transition matrix from Horn (1981), showing the probability of replacement of one individual by another of the same or different species 50 years hence.
Betula populifoliaNyssa sylvaticaAcer rubrumFagus grandifolia
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Using so-called Markov chain model, predicted compositional change over 200 years (and to ∞!)
See initial Betula, then Acer rubrum, then Fagus dominance.
Assumes that transition probabilities from time1 to time2 are constant in space and time and not affected by historical factors such as initial biotic conditions and arrival of species
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Time after disturbance: species invade, then increase, some decrease again and disappear, and some remain as part of the mature vegetation
pioneerspioneers & 'late-invaders'
'Late invaders'
Woody & long lived species
TIME
SEED BANK
Secondary Succession
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In secondary succession after disturbance, two very different kinds of response according to the competitive relationships shown by the species involved.
Founder-controlled – occurs if large number of species are approximately equivalent in their ability to colonise an opening following disturbance, are equally well fitted to the abiotic environment, and can hold their space until they die. Result of disturbance is essentially a LOTTERY. Winner is species that happens to reach and establish itself first.
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Dominance-controlled – occurs when some species are competitively superior (e.g. grow taller, grow faster) to others so that the initial colonisers of an opening do not necessarily maintain their presence there. Result is a reasonably PREDICTIVE SEQUENCE of species because different species have different strategies for exploiting resources. r-selected species are good colonisers and fast growers, whereas later species can tolerate lower resource levels and grow to maturity in presence of early pioneer species and eventually out-compete them.
Secondary succession tends to be a mixture of both kinds of response.
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RESPIRATION
NET PRIMARY PRODUCTION
BIOMASS
TIME
Ecosystem Changes During Succession
1. Changes in biomass and production
PRIMARY SUCCESSION
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RESPIRATION
NET PRIMARY PRODUCTION
BIOMASS
TIME
SECONDARY SUCCESSION
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Primary succession
Small herbsand shrubs
Heath mat
Jack pine,black spruce,
and aspen
Balsam fir,paper birch, &white spruce
climax community
Exposedrocks
Lichensand mosses
Species richness
Biomass
Time
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Secondary succession
Annualweeds
Perennialweeds and
grasses
ShrubsYoung pine forest
Mature oak-hickory forest
Species richness
Time
Biomass
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Biomass accumulation model in secondary succession (102 – 103 years)
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Biomass during stream secondary succession (60 days)
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Soil building during primary succession at Glacier Bay
2. Changes in soil during succession
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Changes in soil properties during primary succession at Glacier Bay
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Changes in soil development nitrogen, pH, cations, organic matter
Nitrogen
pH, cations: Mg & Ca
TIME
Time after fire: secondary succession
Organic matter
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3. Changes in biomass and soil over very long time scales
Hawaiian Islands – volcanic lava flows of different ages extending back to 4.1 million years.
Studied vegetation succession and soil changes, especially soil nitrogen and soil phosphorus.
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Organic carbon and total nitrogen content of soils developing on lava flows Total phosphorus &
percentages of total P in weatherable and refractory (unavailable) forms in soils developing on lava flows
P limitation on oldest soils
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Nitrogen and phosphorus loss rates from soils developing on lava flows
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Biomass changes
Why?
Primary succession
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Recent study on six long chronosequences to investigate reasons for decline in biomass over long time periods.
Wardle et al. 2004 Science 305: 509-513
Birks & Birks 2004 Science 305: 484-485
Six chronosequences Duration (yrs)
Cooloola, Australia Sand dunes >600,000
Arjeplog, Sweden Islands 6,000
Glacier Bay, Alaska Moraines 14,000
Hawaii Lava flows 4,100,000
Franz Josef, New Zealand Moraines >22,000
Waitutu, New Zealand Marine terraces 600,000
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Maxim
al
ph
ase
Retr
og
ress
ive p
hase
Cooloola, Australia
Arjeplog, Sweden
Glacier Bay, Alaska
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Maxim
al
ph
ase
Retr
og
ress
ive p
hase
Hawaii Franz Josef, New Zealand
Waitutu, New Zealand
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Tree basal area – unimodal or decreasing response with age
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Measured C:N, C:P, and N:P ratios for humus and litter
Significant increases in N:P and C:P ratios with age and forest retrogression
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Soil changes:
In the transition from the maximal forest biomass phase to the retrogressive phase, P becomes more limiting relative to N and P concentrations decline in the litter.
N is biologically renewable but P is not, as P is leached and bound in weathered soils.
Over time, P becomes depleted and less available, relative to N.
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Other ecosystem properties:
Also reduced rates of litter decomposition and release of P from litter and decreased activity of microbial decomposers.
Proportion of fungi relative to bacteria increases. Fungal-based food webs retain nutrients better than bacterial-based food webs.
Nutrient cycling thus becomes more closed & essential nutrients, especially P, become less available.
Summary: Long-term decline in biomass is accompanied by increasing P limitation relative to N, reduced rates of P release from decomposing litter, and reductions in litter decomposition, soil respiration, microbial biomass, and ratio of bacterial to fungal biomass.
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(1) changes in species composition and diversity
(2) changes in the structure and function of ecosystems.
What mechanisms drive succession?
Primary and secondary succession in a range of environments and time scales produce
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Mechanisms of Succession
Three mechanistic models – Connell & Slatyer (1977)
1. Facilitation – pioneer species modify environment with time, becomes less suitable for them, and new species invade.
2. Tolerance – initial colonisation by all species, those tolerant of initial conditions become abundant, then species tolerant of new conditions become abundant.
3. Inhibition – initial colonisation by all species, but some species make the environment less suitable for other species, i.e. early arrivals inhibit colonisation by later arrivals.
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Alternative successional mechanisms
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Inhibition of later successional species
Survivorship of successional species under conditions of low tides in hot afternoons
Intertidal successionsSupport for inhibition by Ulva
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Facilitation by algae of colonisation in intertidal succession of surfgrass, Phyllospadix scouleri
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Mt St Helens, Washington. Erupted 1980, created vast new volcanic lava fields.
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Common pioneer plants
1. Anaphalis margaritacea, Epilobium angustifolium – many wind-dispersed small seeds
2. Lupinus lepidus – few large seeds, fixes atmospheric nitrogen
Lupinus lepidus
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Experiments provide evidence for both inhibition and facilitation models
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Lessons from the 25 years of ecological change at Mount St. Helens
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1. Succession is very complex, occurring at different rates along different pathways with periodic setbacks through secondary disturbances (e.g. landslides, mudflows).
2. No single over-arching model of succession provides an adequate framework to explain the observed changes.
3. Chance factors (e.g. timing of the disturbance at various spatial and temporal scales) have strongly influenced survival and successional patterns and pathways.
4. Lakes & most streams largely returned to their pre-1980 state.
5. In contrast, terrestrial vegetation still a mosaic of open areas on steep slopes and eroding sites and well-vegetated areas with shrubs and surviving trees on stable sites.
6. Almost all small mammals have returned but birds have not, possibly because of the lack of extensive forest with vertical structure (niches).
7. Rate of change determined by a complex of factors – position in the landscape, local topography, climate, biotic factors, human factors, and chance.
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Inhibition and facilitation of spruce at Glacier Bay
Primary Succession on Glacial Forelands
Evidence for both inhibition and facilitation
Net I I & F F I effect:
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Are the facilitation, inhibition, and tolerance models useful?
1. Nature is very complex – three mechanistic models are probably a great over-simplification.
2. Real-life situation probably more complex.
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3. General models may not be appropriate for a major ecological process such as succession that consists of a large number of different ecological process – seed arrival, seed bank, competition, herbivory, chance, etc.
C = colonisation
M = maturation
S = senescence
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Despite this undoubted complexity of succession, further mechanisms underlying succession have been proposed
Begon et al. (2006) Chapter 16, pp.483-487
1) Competition-colonisation trade-off and successional niche mechanisms
Early-successional plants have several correlated traits high fecundity
effective dispersalrapid growth rate when resources are abundantpoor growth rate when resources are scarce
Late-successional plants usually have opposite traits
In absence of disturbance, late-successional plants will out- compete early species because they reduce resources (light, water, nutrients) beneath the levels required by early-successional species
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Early species persist because
(1) their dispersal ability and high fecundity permit colonisation and establishment in recently disturbed sites
(2) their rapid growth under resource-rich conditions allows them to out-compete temporarily late-successional species even if they arrive at same time
(1) = competition-colonisation trade-off
(2) = successional niche (early conditions favour early species because of their niche requirements)
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Population density
Population density
temperature
Feeding resource
Feeding resource
temperature
Some Revision!One- and two-dimensional niches
In reality, niche is multi-dimensional
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Realised versus fundamental niche
Fundamental niche = only environment
Realised niche
Biotic control
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Broad and narrow niches
Generalist species
Specialist species
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2) Resource-ratio hypothesis – David Tilman
Rate of changing relative competitive abilities of plant species as conditions slowly change with time.
Species dominance in any point in succession strongly influenced by the relative ability to capture two resources – LIGHT and available SOIL NITROGEN.
Early in succession, the habitat has low N but high light. Nitrogen availability increases with time but light availability decreases with time as biomass increases with time.
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Requirements
Species Light N
A +++ (+)
B +++ +
C ++ ++
D + +++
E + +++
Tilman’s resource-ratio hypothesis of succession
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3) Vital attributes (Noble & Slatyer 1981)
Vital attributes relate to
(1) recovery after disturbance (V = vegetative spread; S = seedling from abundant seedbank in soil; D = dispersal; N = no special dispersal and/or small seedbank)
(2) ability to reproduce in face of competition (T = high tolerance; I = intolerance)
Species then classified on basis of vital attributes
e.g. pioneer Ambrosia artemisiifolia SI
late Fagus grandifolia VT or NT
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4) r and K-selection
Certain attributes are likely to occur together more often than by chance, as expected from an evolutionary perspective.
Two alternatives that increase fitness of a species in a succession
(1) avoids competition, high reproduction, good dispersal, r-selection
(2) tolerant of competition or highly competitive, low reproduction, poor dispersal, K-selection
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r-selection K-selection
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Concept of ‘climax’
Do successions come to an end?
Frederic Clements (1916) single dominant climax in a given climatic region – Monoclimax view
Arthur Tansley (1939) local climax governed by soil, climate, topography, land-use, history, fire – Polyclimax view
Robert Whittaker (1953) - climax-pattern view. Continuum of climax types varying along environmental gradients, not necessarily separable into discrete climaxes.
However, environment is constantly varying at all spatial and temporal scales, so idealised climax is probably never reached in nature, nor is it attainable.
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Community and Ecosystem Stability - Basics
1. Stability – absence of change. May be stable for several reasons (e.g. absence of disturbance, constant environment).
2. In reality, communities and ecosystems are always changing because of changing environment and biotic interactions that may change as organisms age.
3. Stability – ability of community or ecosystem to maintain structure and/or function in the face of potential disturbance.
4. Stability may result from the ability of a community to return to its original state after a disturbance – 'resilience'.
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Succession is the basis for resilience.
Some systems change more quickly than others.
Depends on many factors – climate, soils, available species pool, severity of disturbance, etc.
Require long-term direct observations to study stability and resilience. These are very rare.
Chronosequence is not the same because in the substitution of space for time we assume that the environment has not changed with time.
What Causes Resilience?
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Park Grass Experiment, Rothamsted Experimental Station
Started 1856-1872 to investigate effects of fertiliser treatments on grasslands. Run for over 150 years.
Monitored since 1862.
Shows virtually no new species colonised since 1862.
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1910 – 1948
Three treatments
Proportions changed from year to year (annual rainfall) but relatively stable proportions in the three treatments
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Patterns of species abundance in 60 years
What about individual species?
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Are the Park Grass plots stable or not?
1. Yes, at a very coarse scale – started as a grassland and stayed as a grassland with no new species.
2. Yes, at a less coarse scale of grasses, legumes, and other species but some variation from year to year.
3. No, at the scale of individual species.
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Are there stable natural communities?
Answer dependent on the scale of interest
Environment is changing constantly at a range of scales
Temperature changes in the Northern Hemisphere at different time scales
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Sonoran Desert, Mexico
Saguaro cactus
1959
1984
1998
Repeat photography
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Changes in populations of creosote bushes and saguaro cactus due to major drought in 1960s
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Alternative Stable States and Regime Shifts
Common idea in ecology is of populations and communities fluctuating around some trend or stable average.
Can be an abrupt shift to a dramatically different regime.
Norfolk Broads, England – shallow freshwater lakes showing a rapid regime shift from dominance of aquatic macrophyte plants to a dominance of phytoplankton algae. Regime shift is a result of the use of TBT paint on boats and its toxic effects on gastropod mollusca that graze algae on aquatic plants. (See Lecture 5 Long-term Ecology)
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Saharan desert – gradually declining trend of vegetation cover from 9000 to 5500 years ago, then a sudden collapse into desert.
Changes in sand and silt content in a sediment core near the west African coast
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Coral reefs – very high
biodiversity
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Caribbean coral reefs – sudden dramatic shift of reefs into an algal encrusted state.
Increased nutrient loading as a result of changing land-use promoted algal growth, but this effect did not show as long as herbivorous fish suppressed the algae.
Intensive fishing reduced the fish population and in response the sea-urchin Diadema antilliarum became dominant and became the key herbivore.
When a pathogen killed the dense Diadema sea-urchin population, algae were released from herbivore control, and the coral reefs became overgrown rapidly.
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Different grazers at different spatial scales
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Other examples of dramatic regime shifts:
1. Savannah that is rapidly encroached by shrubs
2. Lakes that shift from clear water to turbid water
3. Standing waters that can suddenly be overgrown by floating plants
4. Different populations in open ocean suddenly change to different abundances synchronously
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Alternate stable states – How can they occur?
Although plants compete for resources, this competition can be overruled by facilitation because the vegetation ameliorates certain critical conditions.
Terrestrial vegetation in dry regions can enhance soil moisture and microclimatic conditions.
Leads to positive feedback between vegetation and moisture
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1. Precipitation in absence of vegetation is determined by climate
2. Vegetation has a positive feedback on local rainfall
3. No vegetation when precipitation falls below critical level
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Actual precipitation can be drawn as two different functions of global climate; one without vegetation, one with vegetation.
Above the critical level, vegetation is present. Below the critical level, vegetation is absent.
If general climate gets wetter, only the plant regime exists. If very dry, regime of no vegetation.
Over a range of climatic conditions, two alternative stable states or regimes can exist.
Instability between Fc and Fd
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Shallow freshwater lakes and two alternative stable states
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Stability landscapes showing resilience of equilibria
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Ball (state of ecosystem) tends to settle in 'valleys' = stable regime state.
'Hill' between the 'valleys' is barrier between two alternative states or regimes.
Changes in external conditions can change the stability landscape by changing the depth of the 'valleys' and the height of the 'hill'.
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Nutrient level
Algae-dominated
Plant-dominated
Macrophyte-dominated system
pre-1960
Use of TBT in boat paints
1960
Plant-dominated
state
Decrease of mollusca (gastropods, etc.)
Increase in algae
Decline of macrophytes
Reduction in grazing of
epiphytic algae
Algae-dominated
state
See Lecture 5 Long-term Ecology for details
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1 & 4 - alternate states, 2 - causes of change3 - triggers of resilience and regime shifts
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Reduced resilience makes the system vulnerable to a regime shift
Resilience of the low P input state is high as the likelihood of crossing the threshold from one state to another is low (big distance between the two states).
Resilience of the high P input system is low as the likelihood of crossing the threshold from one state to another is high (low distance between the two states).
(a)
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Evidence from field data
(a) Pacific Ocean
(b) Dutch ditches
(c) Shade in shallow lakes
= dominated by cyanobacteria
= dominated by other algae
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Alternative stable states – can they be predicted?
Beaugrand et al. 2008 Ecol Letters 11: 1157-1168
North Atlantic – critical thermal boundary where a small increase in temperature triggers abrupt ecosystem shifts across multiple trophic levels.
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Boundary is located where abrupt shifts occur.
All closely related to annual sea-surface temperature (SST).
Critical at 9-10°C, establishment of Westerly winds marine system.
Beaugrand et al. 2008
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Decadal changes in SST 1960-2005 and predicted changes in 2090-2100
Small changes in last 40 years
Beaugrand et al. 2008
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Ecosystem state shifts between 1986 and 1988, preceded by a period of high ecosystem variability
Pre-1981, 72% of cells have SST of 9-10°C; post-1988, 20%
Major shift in SST affecting many aspects of ecosystem. Shift predicted by increasing variance in biological systems
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What of the future?
Two future climate scenarios: progressive shift northwards from 2000 to 2090
Climate changes in SST will alter biodiversity and carrying capacity of ecosystems.
Changes will precipitate major reduction in stocks of Atlantic cod, already severely impacted by exploitation from fishing.
Relatively small climatic change may ‘tip the balance’ in an already over-exploited ecosystem (reduced resilience)
Beaugrand et al. 2008
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Summary of Alternative Stable States
1. System has alternative states if there can be more than one 'stable state' for the same external variable (e.g. nutrients in lakes).
2. Stable states are really dynamic regimes. Show slow trends, natural population fluctuations due to climate and internal population dynamics.
3.Multiple causes are the rule in regime shifts.
4. Patterns depend on spatial scale. May have a mosaic of alternative stable states. May remain unaltered until an extreme event triggers a shift in the patterns.
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5. External conditions should really be external and independent and not an interactive part of the system.
6. External conditions may be affected by the system if the change in external conditions is very slow relative to the natural rates of change in the system.
Collapse of vegetation in the Sahara occurred over 100-200 years but this is fast compared with the forcing function, namely gradual changes in the Earth's orbit.
7. In some systems, fast and slow components can affect each other mutually and this leads to population cycles (e.g. recurrent pest outbreaks).
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8.Resilience is necessary to sustain desirable ecosystem states in variable environments and uncertain futures.
9. Humanity has drastically altered the capacity of ecosystems to withstand or 'buffer' disturbance. Cannot assume that there will be a sustained flow of ecosystem 'services' or functions to our well-being.
10.Biological diversity appears to enhance the resilience of ecosystem states
11."Nature is not fragile … what is fragile are the ecosystem 'services' on which humans depend" Simon Levin (1999)
What causes natural population fluctuations, the fluctuations around some mean in one 'stable state'?
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Maintenance Dynamics
Even if the environment is stable (which it never is!), there are factors INTERNAL to the community that cause change, so-called 'cyclic' succession.
Cycle of events replicated many times over the whole of the community as a series of PHASES. Provides a mosaic of phases within community. PATCH DYNAMICS
Succession is a directional change
Cyclic changes or maintenance dynamics or patch dynamics are fluctuations about a mean value.
A.S. Watt ‘Pattern and Process’ 1947Dr Alex ‘Sandy’ Watt
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Phases in plant growth with age
pioneer
building
mature degenerate
age
pro
du
ctiv
ity
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Phases in growth of Festuca ovina
Changes in cover of three species 1936-1973 (F. = Festuca, H. = Hieracium, T. = Thymus)
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Important factors in maintenance or patch dynamics
1. Disturbance (or ageing) gaps
2. Dispersal recruitment growth
3. Frequency of gap formation
4. Size and shape of gaps
View landscape as patchy with disturbance and recolonisation by individuals of different species
Critical roles for disturbance (and ageing) as a RESET mechanism, for dispersal and establishment between habitat patches, and competition between species concerned
Community dynamics need a landscape-scale perspective to be understandable
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Fire: control of secondary succession in west Norwegian coastal heathlands
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Time
Calluna spirer
+ urter og gress
forveet calluna
trær
Bjørk og fufu skog ( Eik)
FIRE!BRANN!
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Fire also important in community maintenance dynamics – fine-scale burning
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Burnt versus unburnt heath
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Mosaic of burning phases
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Maintenance dynamics of Calluna (røsslyng) in coastal heathlands involving fire
Traditional heathland cycle
Dereliction
pioneer
building
mature
degenerate
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Combination of controlled burning, mowing, & grazing
'Cultural landscape'
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Disturbance and Diversity
Disturbance resets the clock in any succession. Elimination of existing populations, allows colonisation by early successional species - frequency of disturbance critical.
a)high frequency of disturbance, pioneers only
b)intermediate disturbance, pioneers plus later species, giving maximum diversity
c) low disturbance, late species only
Result is hump-backed curve of diversity in relation to disturbance 'intermediate disturbance hypothesis'
Hypothesis formulated in relation to successional responses after disturbance.
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Community Concepts Revisited
Palaeoecology – study of the distribution & abundance of organisms (plants and animals) in the past.
Pollen analysis – major technique.
Last glaciation about 18000 years ago and subsequent deglaciation at about 11000 years ago were a major, broad-scale primary succession.
Extent of glacial ice at 18000 and 8000 years ago
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Large number of sites where pollen analysis has been done. Can determine when a particular tree arrived and expanded at a site and then map the times of tree arrival to detect tree migration patterns since the last deglaciation.
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Each tree genus has its own individualistic history. Did not move as forest communities.
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Same in the British Isles – strongly individualistic behaviour of forest trees
Bjørk Hassel
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Eik Alm
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Furu Lind
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Organismal concept – F.E. Clements
Individualistic concept – H.A. Gleason
In fact these two concepts refer to different scales and biological concepts – no real conflict!
Organismal concept is a spatial concept
Individualistic concept is a population concept
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4 species populations along an environmental gradient (vertical plot)
4 species along a geographical or spatial gradient (horizontal gradient)
Can recognise several communities along spatial gradient – A, A+B, B, C, D, and transitions B+C and C+D
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Great Smoky Mountains, Eastern USA
Robert H. Whittaker
Landscape distribution of vegetation types
Spatial arrangement of vegetation types
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Environmental distribution of populations – individualistic concept
Landscape or spatial distribution of vegetation
types – organismal concept
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Community structure is thus the product of a complex interaction of pattern and process in space and time.
Each species responds to a wide range of environmental factors that vary continuously in space and time across the landscape.
Interactions between organisms influence the nature of these responses.
The end result is a dynamic mosaic of communities within the landscape.
Study of this mosaic at the landscape scale is landscape ecology (see Lecture 7 on Landscape and Geographical Ecology).
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Conclusions and Summary
1. Succession is the gradual, directional change in plant and animal communities in an area following the creation of new substrates (primary succession) or disturbance (secondary succession).
2. Succession generally ends with a mature community that is similar to the surrounding vegetation and fauna and that has relatively stable populations ('climax' vegetation).
3. Environment varies at a wide range of temporal and spatial scales.
4. Primary succession has been studied in detail on glacial forelands in western North America and Norway. Moraines of different but known ages provide a chronosequence.
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5. Plant abundance, species composition, and species richness change over time. Richness increases and then often declines with time.
6. Ecosystem changes during succession include increases in biomass, primary production, soil composition, and nutrient retention. Phosphorus limitation becomes more important in 'old' systems.
7. Mechanisms to explain succession include facilitation, tolerance, and inhibition.
8. Field evidence provides support for facilitation, inhibition, or a combination of the two.
9. Nature is more complex than 3 mechanistic models. Succession is a combination of many different ecological processes –germination, herbivory, competition, chance, etc.
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10. Community stability may be due to a lack of disturbance or community resistance ('resilience') to disturbance.
11. Communities are both stable and unstable, depending on scales of study. Alternative states can exist and catastrophic regime shifts can occur.
12. Within-community maintenance dynamics or patch dynamics ('cyclic' changes) are what makes a community maintain itself.
13. Human activity can prevent secondary succession and can influence maintenance dynamics, to create so-called cultural landscapes.
14. Succession occurs over a wide range of time scales ranging from days, months, centuries, to millions of years. Basis of ecological change.
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15. Palaeoecological data indicate that forest trees showed individualistic behaviour in their migration patterns after the last deglaciation.
16. The community is a spatial concept. The individualistic continuum is a population concept.
17. The real world lies between the organismal and individualistic concepts, depending on our spatial and temporal scales of study and on our choice of gradient (spatial, environmental).
18. Vegetation at the landscape scale is a mosaic depending on topography, environment, primary succession, secondary succession, and maintenance dynamics.
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EECRG Research Topics in this Lecture
www.eecrg.uib.no
Primary succession on glacial forelands in Norway, Nepal, and Tibet
Alternative stable states in Norwegian forest vegetation
Natural climatic variability in NW Europe in the last 15000 years
Tree migration patterns in the last 12000 years
Ordination gradient analysis of many different vegetational and faunal communities
Heathland ecology, management, and dynamics in western Norway