Community formation

and composition
Martin Sullivan
martin.sullivan@mmu.ac.uk
JD E413
Formation of communities
• Mount St Helens eruption
destroyed existing
vegetation
• Blank canvas for new
communities to form
Formation of communities
• Mount St Helens eruption
destroyed existing
vegetation
• Blank canvas for new
communities to form
• Succession – the process
of community assembly
and development

Virginia Dale studied succession

following the eruption by monitoring
permanent plots through time
How do communities form (succession)?

Why do we get different communities

in different places?
Some definitions - succession
Succession: Changes in species composition and
abundance over time as communities develop
following disturbance
Primary succession if the disturbance has killed
almost all organisms (or the habitat is completely
new, e.g. a new volcanic island)
Secondary succession if the disturbance is more
minor and only some organisms are killed
Some definitions - community
Community: A group of populations that interact
with each other
Spatial community: All organisms that occur in a
particular locality
Can be refined to be just species of a particular
trophic level (i.e. primary producers or herbivores –
trophic community), just species that use a
particular resource (guild – e.g. seed-eating birds), or
just species in a particular taxonomic group (e.g. just
mammals – taxonomic community).
A lot of work has focused on plants as they are
easy to study, but concepts apply to microbes and
animals

Microbes
• Antibiotics act as a disturbance agent, killing many
microbial organisms
• Consequences of this disturbance for microbial
community important for our health
A lot of work has focused on plants as they are
easy to study, but concepts apply to microbes and
animals

Animals
• Bird communities change as forest plantations age
• Animal communities are to some extent passengers
of change in plant communities

Tree pipit in young plantation Common crossbill in mature plantation

 Which species can colonise
following disturbance?
Source pool – set of species that can
potential disperse to and reach the
site
 Environmental filters determine
which species can colonise
Source pool

Environmental filter –
which species can
tolerate the
environmental
conditions at a location?

Community
Environmental filtering relates to
species’ niches
• Niche – the set of environmental conditions in
which a species can survive, grow and reproduce
Precipitation

Temperature
 Environmental filtering relates to
species’ niches
• Niche – the set of environmental conditions in
which a species can survive, grow and reproduce

Favourable biotic
Abiotic
environments
[environmental]
(e.g. prey species,
conditions a species
lack of
Realised can survive in
competition)
niche Fundamental
niche

Areas a species can

potentially move to
Environmental filters mean that it is possible to
predict environmental conditions from species
composition

• Heinz Ellenburg developed a set of indicators that can be

used to predict environmental conditions (e.g. light,
moisture)
• Created a database where each species has a value that
reflects their environmental affiliation
• Averaging this across a community gives a good
approximation of what the environmental conditions
actually are
 The species that colonise first can alter the
environment to make it more favourable for other
species - facilitation

Facilitated succession described

by Connell & Slayter 1977,
American Naturalist
Facilitation in saltmarsh development
 Some plants ecosystem engineers
 Trap sediment and increase elevation
 Increase soil oxygenation
 Allow other plants to colonise – aka facilitated succession
Sediment accretion associated with Spartina

Spartina anglica

Rosso et al. (2006) Rem Sens Env, 100, 295-306

See also Castellanos et al (1994) "Nucleation and facilitation in saltmarsh succession: interactions between Spartina maritima and
Arthrocnemum perenne." J. Ecol 82, 239-248.
Sedimentation makes saltmarsh
 Tides bring in sediment
 Frequent inundation = more sedimentation
 So as marsh gets higher = fewer inundations = less
sedimentation
 They can keep pace with sea level….at the moment…
Highest tides
sedimentation
saltmarsh

Low water Springs

mudflat

Sea Land
 Mudflat
Low marsh Competitive
This leads to ability
Flooding
the zonation tolerance

we observe Mid marsh

High marsh
Succession and saltmarsh creation
• New saltmarshes created
by managed realignment
to compensate for losses
• We expected them to
develop in the same way
as natural marshes
• But we found that the
communities that
developed differed from
those on natural marshes
 Why did created saltmarshes develop
differently?

Sullivan et al. 2018 Journal of Applied Ecology

 In created saltmarshes, who gets there
first wins
• Plants can colonise bare earth across the elevations
• So speed of dispersal likely to matter a lot
• Who gets there first can dominate

Sullivan et al. 2018 Journal of Applied Ecology

 This is an example of inhibition
succession
• Under Connell and Slayter (1977)’s model of
succession, inhibition succession can occur when the
species that get there first inhibit the establishment
of other species
• Creates a big problem in ecological restoration –
chance events can dictate the development of a site
Different strategies for species to win at
succession
• J. Phillip Grime came up with the CSR scheme for classifying species
• Originally for plants, but potentially applicable to all organisms
• Classified species as:
• Ruderal – I disperse really well and will get there first after
disturbance
• Stress – I can cope with really tough environmental conditions
• Competition – I don’t care if you are already there, I’ll
outcompete you
• Trade-offs prevent species being good at
everything C

R S
Different strategies for species to win at
succession

Ruderal Competition

Stress Competition

R S
Climax community – the endpoint of
succession
• The stable community that forms following succession (until
the next disturbance)
• Shaped by environmental conditions at broad and local
scales

WWF terrestrial ecoregions Soil fertility determines whether you get dry
forest or savannah in parts of Brazil
Climax community – the endpoint of
succession
• Small disturbances (e.g. tree-fall events) create micro-
succession within the climax community
• These disturbance events can be important for allowing
species (e.g. ruderal species) to persist
 Succession is often thought of as a
predictable process, but outcomes can be
influenced by chance events
Back to Mount St Helens
• Nutrients washed into Spirit Lake
• Switched bacterial community to be
dominated by chemosynthetic taxa
(opposed to photosynthetic)
• Change came a few years later when
heavy rains diluted nutrients
• Phyto and zooplankton communities then
rapidly recovered
• But fish still dispersal limited – chance
introduction of rainbow trout
 Succession is often thought of as a
predictable process, but outcomes can be
influenced by chance events

• Seeding with
grasses can
inhibit
establishment of
forest

See Stuble et al. 2017 J Applied

Ecology for an example of how
chance affects ecological restoration
https://besjournals.onlinelibrary.wile
y.com/doi/10.1111/1365-2664.1286
1
Conservation often seeks to maintain
early successional habitats
• Bitterns live in reedbeds – early successional habitat that dries
out and turns into woodland
• Population collapsing towards local extinction in the UK
Conservation often seeks to maintain
early successional habitats
• They need wet (v early successional) reedbeds (see Gilbert et al.
2005 Ibis)
• Increasing disturbance creates more of this habitat – now
common practice in nature reserves
Conservation often seeks to maintain
early successional habitats
• Stone curlew another declining bird in the UK

• Needs open ground for nesting (to see predators), but some
tussocky vegetation to find insects to eat

• This habitat relies on regular disturbance

• Species does well in military firing areas – activities create

necessary disturbance regime
 Recap
• Communities develop by succession until they reach the climax
community
• The species that can colonise a location have to be able to get
there (source pool) and survive there (environmental filtering/
the niche)
• Facilitation by early colonists can make environmental conditions
more favourable for later establishing species
• But early colonists can also inhibit species establishment
• Species can be classified into competitive, stress tolerant or
ruderal strategies
• Chance events can alter the outcome of succession – makes it
harder to restore habitats
• Conservationists often fight succession to maintain valuable
early-successional habitats