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Transcript
Community Ecology
Biosphere
Chapter 54
Population
Ecosystem
Organism
Community
1
Community
•
2
Biological communities
A biological community refers to all the
species that occur together at any
particular locality.
3
4
Gleason & Clements
Individualistic
Holistic
Individualistic
Holistic
5
6
1
Number of stems per hectare
Individual species in a community respond
independently to changing environmental conditions.
A synchronous change of species in communities
exist only when sharp changes of environmental
conditions occur.
Normal
soil
400
Ecotone
Serpentine
soil
Black oak
Poison oak
Iris
Douglas fir
Hawkweed
California fescue
Snakeroot
200
Canyon live oak
Collomia
Ragwort
Yarrow
Buck brush
0
Small fescue
Fireweed
Knotweed
Dry
Wet
Moisture gradient
7
Biodiversity: number & evenness
Attributes of ecological communities
–
–
–
8
Biodiversity —
ƒ The number and evenness of species.
kingfisher
merganser
otter
great blue heron
dipper
garter snake
The niche concept—
ƒ The role of interspecific interactions
steelhead
roach
stickleback
water scavenger frog tadpole
beetle larva
Food web structure —
ƒ The feeding relationships among species.
newt
caddis fly larva
snail
crayfish
tuft midge
–
Succession—
ƒ The change of communities through time.
diatoms
9
Biodiversity measures the number of
species and their evenness
•
blue-green algae
green algae
10
Most species in a community are rare
Species
richness
may be equal,
but relative
abundance
may be
different.
11
12
2
Exercise: What factors determine the
occurrence of a species in a community?
Biodiversity: community 1 > 2
kingfisher
merganser
great blue heron
otter
dipper
garter snake
steelhead
roach
stickleback
water scavenger frog tadpole
beetle larva
snail
newt
caddis fly larva
crayfish
tuft midge
diatoms
13
First, a species needs to be able to tolerate
the local physical-chemical environment.
green algae
blue-green algae
14
Second, the environment must provide
required resources, such as food.
+ required resources
15
16
A niche
~ the fundamental “niche”
+ required resources
17
18
3
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
The distribution patterns
of Chthamalus & Semibalanus
Joseph Grinnell (UC Berkeley)
High tide
Chthamalus
Low tide
Semibalanus
19
20
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Exercise: Is this range the fundamental niche of
Chthamalus? How do you know?
Joseph H. Connell (UC Santa Barbara)
High tide
Chthamalus
Low tide
Semibalanus
21
Results: What does the removal experiment tell you?
Let’s do a removal experiment
High tide
[1] removing
Chthamalus …
High tide
Chthamalus
After removing
Chthamalus …
Chthamalus
Low tide
Semibalanus
22
Low tide
[2] removing
Semibalanus …
23
Semibalanus
After removing
Semibalanus …
24
4
Fundamental vs Realized Niches
Realized Niches
–
High tide
Actual niche utilized by an organism is
influenced by competition, predation,
parasitism, mutualism, etc..
Chthamalus
Low tide
–
Biological interactions change niche space,
thus contribute to the structure of a
community.
Semibalanus
Fundamental Realized
niches
niches
25
Predation often reduces “niche” space
~ the realized “niche”
26
Biotic interactions limiting
+ required resources
27
28
Exercise: What’d happen if 2 species had
overlap niches, e.g. steelhead & roach?
Ghost of competition past …
Ghost of predation past …
The fundamental niche of a species may shift
due to consistent biological interactions over
evolutionary time scale. Thus, the effects of
competition or predation are no longer exist in
the present day.
kingfisher
merganser
great blue heron
otter
dipper
garter snake
steelhead
Species as we see them today are products of
biological interactions over evolutionary times.
roach
water scavenger frog tadpole
beetle larva
snail
newt
caddis fly larva
crayfish
tuft midge
diatoms
29
stickleback
green algae
blue-green algae
30
5
Outcomes of interspecific competition
–
One species may be driven to extinction (competitive
exclusion).
When resources are limiting, no two
species utilizing the same niche can
coexist indefinitely.
– one will eventually eliminate the other
Or, one or both species has to shift their niches by
changing their behavior (resource partitioning).
Sometimes, morphology change as well (character
displacement) to enhance resource partitioning.
31
32
Competitive Exclusion Among Paramecium
Georgyi F. Gause (Moscow Univ.)
Alone
Population density
–
•
200
200
150
P. aurelia
Together
100
50
50
0
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
P. caudatum
P. aurelia
150
100
50
0
50
0
4
8 12
Days
16
20
24
34
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
200
150
200
150
100
0 4 8 12 16 20 24
Days
Either way, competition reduce population sizes, thus,
change relative abundance and community structure.
If competitive exclusion did not occur,
resource partitioning must exist.
Alone
P. caudatum
0
0 4 8 12 16 20 24
Days
0
33
150
100
200
Population density
–
If niche overlap extensively, then, to coexist, the shared
resources must not be limiting. Otherwise, …..
P. caudatum 100
P. bursaria
50
0
0 4 8 12 16 20 24
0 4 8 12 16 20 24
Days
Days
P. caudatum
P. bursaria
Together
Population density
–
Principle of Competitive Exclusion
75
Eat bacteria
50
Eat yeasts
25
0
0
4
8
12
Days
16
20
35
36
6
Consequences of long-term competition
•
Resource Partitioning
The fundamental niche of a species may shift
under long-term interspecific competition
through 2 processes:
–
Resource partitioning
–
Character displacement
•
Species living in the same area (sympatric
species), over time, evolve behavioral
mechanisms that partition available
resources to avoid direct competition.
37
38
Resource Partitioning – the sympatric species
have subdivided the niche
Robert MacArthur
(Princeton Univ.)
RESOURCE PARTITIONING OF INSECT-EATING WARBLERS
Cape May
warbler
Bay-breasted
warbler
Myrtle
warbler
39
40
Resource partitioning of insect-eating lizards
Character displacement
•
41
Sympatric species tend to exhibit greater
differences in morphology than allopatric
species (species that live apart from each
other) to avoid direct competition.
42
7
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Does the diversification of Darwin’s finches involve
resource partitioning & character displacement?
Individuals in each size class (%)
Character Displacement
50
G. fuliginosa
Alone
Los Hermanos
Islets
25
0
50
Daphne Major
Island
25
0
50
G. fortis
Alone
San Cristobal and
Floveana Islands
25
0
G. fuliginosa
and
G. fortis
Sympatric
7
9
11 13
15
Finch beak depth (mm)
43
44
Exercise: Use an analogy to describe “niche”
to your friend.
Mode of interspecific competition
•
kingfisher
merganser
otter
great blue heron
dipper
garter snake
steelhead
roach
stickleback
water scavenger frog tadpole
beetle larva
snail
newt
•
caddis fly larva
crayfish
tuft midge
diatoms
green algae
Exploitative competition
– e.g. Paramecium compete for food
– the 2 species consume shared resources
– The 2 species don’t need to see each other
Interference competition
– e.g. barnacles compete for space
– the 2 species fight over resources (space)
– there are face-to-face confrontations
blue-green algae
45
Exploitative competition – Hey, don’t drink too fast !
46
Interference competition – Hey, you pinch my straw !
47
48
8
Kangaroo rats and other desert rodents
Number of other rodents
Removal experiments are the strongest tests of
the existence of interspecific competition
Kangaroo rats removed
Kangaroo rats present
15
10
5
0
1988
1989
1990
49
Exercise: So, how does competition affect
community structure?
Detecting Interspecific Competition
•
50
Negative effects of one species on another
do not automatically indicate competition.
kingfisher
–
Presence of one species may attract a
predator that consumes both, causing one
species to have a lower population size
than the other. (apparent competition)
merganser
great blue heron
otter
dipper
garter snake
steelhead
roach
stickleback
water scavenger frog tadpole
beetle larva
newt
snail
caddis fly larva
crayfish
tuft midge
diatoms
51
Predation also affects community structure.
kingfisher
merganser
green algae
blue-green algae
52
Exercise: What’d happen to the
community when heron reduce roach?
otter
great blue heron
kingfisher
dipper
garter snake
merganser
great blue heron
otter
dipper
garter snake
steelhead
roach
stickleback
water scavenger frog tadpole
beetle larva
newt
steelhead
caddis fly larva
snail
roach
stickleback
water scavenger frog tadpole
beetle larva
crayfish
tuft midge
snail
newt
caddis fly larva
crayfish
tuft midge
diatoms
green algae
blue-green algae
diatoms
53
green algae
blue-green algae
54
9
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Number of individuals
Predators reduce the population size of prey, and
often induce top-down trophic cascades.
Trophic cascades are a
key feature in
community dynamics.
Paramecium
1200
We will talk about it
in ecosystem ecology.
800
Didinium
400
0
1
2
3
4
5
6
7
Days
55
Coevolution
56
Plant defenses against herbivores
-- Morphological defenses
The interdependent evolution of two or more
species.
Particularly, “arms races” between predators
and prey drive evolutionary changes and
remarkable adaptations in both predator and
prey species, and promote species diversity.
57
Plant defenses against herbivores
-- secondary compounds
58
Some herbivores can breakdown
secondary compounds
Mustard oils – cabbages – cabbage butterfly
59
60
10
Herbivores make use of
secondary compounds
Herbivores make use of
secondary compounds
Cardiac glycoside – milkweed – Monarch butterfly
61
Chemical defenses
Prey defenses against predators
•
Chemical defenses
•
Morphological defenses
–
Cryptic (camouflage) coloration.
–
Warning (aposematic) coloration.
–
Mimicry
62
63
Camouflage! Can you see them?
64
Warning! Dare you eat them!
65
66
11
Batesian mimicry is where a harmless
species mimics a harmful one.
Danaus plexippus
Müllerian mimicry is where two or more
unpalatable species resemble each other.
Limenitis archippus
67
68
In addition to competition, predation & herbivory,
there are other types of species interactions
Müllerian mimicry & Batesian mimicry
69
70
Commensalism
Cannibalism
Epiphytes
71
Barnacles on whales
72
12
Mutualism
Parasitism -- ectoparasites
73
Parasitism -- endoparasites
74
Insect Parasitoid
75
Parasites with complex life cycles often
manipulate host behavior
76
Mutualism turns parasitism
77
78
13
What’d happen to ants
if you removed rodents?
Interactions Among Ecological Processes
•
•
Predation may reduce competition
Parasitism may alter competition
Indirect effects
– Presence of one species may affect a
second species through interactions with a
third species.
–
Ants
Rodents
+
+
–
–
Large seeds
79
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
60
Small seeds
80
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
ComplexArt)
indirect interaction
Fig. 54.23(TE
Fig. 54.22(TE Art)
Number of ant colonies
•
Rodents removed
Rodents not removed
–
Ants
40
Rodents
20
–
+
+
–
Oct 74 May 75 Sep 75 May 76 Aug 76 Jul 77
Sampling periods
Large seeds
Small seeds
81
Keystone species
82
A tidal pool community
A species whose absence would
bring about a significant change in
the community.
ƒ Top predators in a community are
often keystone species (e.g. sea
stars affect the diversity of tidal pool
communities).
ƒ
83
84
14
After sea star removal
Before
After
85
86
Exercise: Who do you think is the
keystone species in this community?
Sea stars as a
keystone species
kingfisher
merganser
great blue heron
otter
dipper
garter snake
steelhead
roach
stickleback
water scavenger frog tadpole
beetle larva
newt
snail
caddis fly larva
crayfish
tuft midge
diatoms
87
Ecosystem engineering
green algae
blue-green algae
88
Succession
Some keystone species modify their habitats to
produce a particularly strong effect on community
structure, e.g. beavers.
•
Biological communities change with time,
usually, but not necessarily, from a simple to
a more complex structure.
–
–
89
primary succession
secondary succession
90
15
Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.
Soil nutrients change as succession progress
Primary succession - occurs on bare substrates
300
Year 1
Glacial retreat
Nitrogen (g/m2)
250
Year 100
Year 200
c
200
Nitrogen
in forest
floor
150
100
b
50
a
Year 1
Pioneer
mosses
91
Nitrogen
in mineral
soil
Year 100
Year 200
Invading Alder
Spruce
alders
thickets forest
Secondary succession occurs where an existing
community has been cleared, but the soil is left intact.
Primary succession - occurs on bare substrates
93
94
Why succession occurs?
•
•
•
92
Why succession occurs?
Pioneer species are able to tolerate harsh
conditions during early succession.
•
Seed dispersers also aid the succession
process
Settled species alter the habitat which
facilitate the invasion of subsequent
colonizers.
The habitat may be further altered that inhibit
the growth of original colonizers.
95
96
16
Marine and island communities are subject
to disturbance by tropical storms.
Most communities are in a state of nonequilibrium
owing to frequent disturbances
97
We often think that disturbances have a
negative impact on communities, but some
communities are maintained by disturbance.
98
Species number
Exercise: Do disturbances increase or decrease
species diversity in a community? How?
?
Disturbance
99
Exercise: What factors influence the
biodiversity of this community?
Intermediate disturbance theory
ƒ
100
Communities experiencing moderate
levels of disturbance will have
greater species richness than
communities experiencing either
smaller or larger levels of
disturbance.
™ simultaneous successional stages
™ dominant competitors kept at bay
kingfisher
merganser
great blue heron
dipper
garter snake
steelhead
roach
stickleback
water scavenger frog tadpole
beetle larva
snail
newt
caddis fly larva
crayfish
tuft midge
diatoms
101
otter
green algae
blue-green algae
102
17
Causes of biodiversity, so far
ƒ
ƒ
ƒ
ƒ
Biosphere
Physical-chemical environment
Biological interaction
Disturbance
Others?
Population
Ecosystem
Organism
Community
103
104
Dynamics of Ecosystems
Ecosystem ecology
Chapter 55
•
•
Chapter 55
An ecosystem includes all the organisms
living in a particular place, and the abiotic
environment in which they interact.
Ecosystem ecology is about …
– Biogeochemical cycles of chemical
elements through ecosystems.
– Energy flows through ecosystems (a
one-way process).
105
Eugene Odum (Univ. of Georgia)
106
Biogeochemical Cycles
•
•
•
•
107
The water cycle
The carbon cycle
The nitrogen cycle
The phosphorus
cycle
108
18
Physical processes move elements around
Biological processes move elements around
Feeding
Photosynthesis
Decomposition
109
110
The water cycle
Biogeochemical Cycles
•
•
•
•
The water cycle
The carbon cycle
The nitrogen cycle
The phosphorus
cycle
Ice
Organisms
111
112
Reservoirs of water
The Water Cycle
•
•
Atmosphere
By area, oceans receive ¾ and lands ¼ of
precipitation. Both lands and oceans
evaporate much of water back to the
atmosphere.
Ice
On lands, transpiration from plants account
for 90% of water back to the atmosphere.
Organisms
113
114
19
Water cycling processes
Groundwater
•
Ice
•
Organisms
Groundwater (stored in aquifer) is the largest
reservoir of water on lands.
It is recharged via percolation of rainfall and
water seeps down from lakes and streams
through soil, very slowly. The flow is also
very slow.
115
116
Mining and pollution of groundwater
•
•
Deforestation
breaks the local water cycle
•
Human mine groundwater, thus many
underground aquifers have much higher
withdraw rates than recharge rates.
Agriculture will be in trouble.
Pesticides, herbicides, and fertilizers
accumulate in groundwater are virtually
impossible to be removed (b/c slow
turnover rate). Yet, groundwater supplies
about 50% of drinking water (in U.S.).
•
In forest ecosystems, plants take up much
moisture and transpire it back into the
atmosphere, forming clouds. When forests
are cut down, water drains from the local
area instead of forming clouds.
The lack of rains further prevents
reforestation, and may create semi arid
desert (positive feedback).
117
Deforestation also causes erosions
118
Biogeochemical Cycles
•
•
•
•
119
The water cycle
The carbon cycle
The nitrogen cycle
The phosphorus
cycle
120
20
The carbon cycle
Carbon Cycle
•
•
Carbon cycle is based on CO2 which
makes up 0.03% of the atmosphere,
dissolved in water, and those C locked up
in organic compounds (live or dead
organisms) and sediment (carbonates).
Most organic compounds formed as a
result of CO2 fixation (photosynthesis).
About 70% of CO2 in the atmosphere is
fixed by photosynthesis annually.
121
122
Reservoirs of Carbon
Carbon Cycle
•
•
Carbon in the ecosystems cycles through
photosynthesis and respiration processes.
Most organic compounds are ultimately
broken down, and C is released back into
the atmosphere.
Under natural condition, photosynthesis
and respiration are proximately balanced.
But we have been tipping the balance.
123
124
Carbon cycling processes
Tipping the balance
•
125
Increasing fuel
consumption, and
deforestation by
burning are
liberating carbon
at an increasing
rate.
126
21
Green house effects
CO2 and global warming
127
128
The nitrogen cycle
Biogeochemical Cycles
•
•
•
•
The water cycle
The carbon cycle
The nitrogen cycle
The phosphorus
cycle
129
130
Nitrogen-fixing symbionts
The Nitrogen Cycle I
•
•
Only a few kind of prokaryotes can fix
atmospheric nitrogen into forms that
can be used for biological processes.
Only symbiotic bacteria fix enough
nitrogen to be of major significance
Nitrogen fixation: N2+ 3H2 Æ 2NH3
•
•
131
Nitrogen fixation:
N2+ 3H2 Æ 2NH3
Nitrogen-fixing
bacteria form
symbiotic
relationship
mainly with the
roots of legumes
(the pea family).
132
22
Reservoirs of nitrogen
The Nitrogen Cycle II
•
After organisms die, nitrogen in organisms
become soil nitrates through 2 processes
carried out by decomposers.
Organisms
•
•
Ammonification – NH4+ (ammonium) are
released during the metabolic processes of
decomposers.
Nitrification -- NH4+ are converted by bacteria
into NO2- (nitrite) and NO3- (nitrate) which can
be taken up by regular plants.
133
Nitrogen cycling processes
The Nitrogen Cycle III
•
•
134
Nitrogen fixed in soil (NO3- , nitrate)
may return to atmosphere through
Denitrification, carried out by bacteria
under anaerobic condition.
Feeding
Denitrification: Bacteria get O2 from
NO3- and release N2 or N2O back to
the atmosphere.
135
Tipping the balance
•
136
Eutrophication
The use of
fertilizers, thus
large amount of
nitrogen, in
agriculture alter
the natural cycles
of nitrogen.
137
138
23
Phosphorus Cycle
Biogeochemical Cycles
•
•
•
•
The water cycle
The carbon cycle
The nitrogen cycle
The phosphorus
cycle
139
The Phosphorus Cycle
•
•
140
Reservoirs of phosphate
Organisms
The largest reservoir of phosphates exist
in rocks.
Phosphates weather from rocks and soils
into water. They may enter plants and
animals, or are transported to oceans and
accumulate in sediments.
141
Sediments & up-welling
•
142
Guano deposits
The nitrates and phosphates locked up in
sediments, particularly those in the deep
ocean floor, can be brought back only by
up-welling.
143
144
24
Tipping the balance
•
Eutrophication
Three times more
phosphates than
crops required are
added to
agricultural lands
each year.
145
146
Exercise: What would happen to nutrient
cycles after deforestation?
When the watershed is intact …
The amounts of nutrient inputs from rains
and snows are about equal to the amounts
that flow out.
•
When the forest is cut down …
•
Hubbard Brook Experimental Forest
The nutrient inputs from rains drain from the
local area.
147
Consequences of deforestation
Nutrients cycle,
but energy doesn’t
80
Concentration
of nitrate (mg/l )
148
40
4
Deforestation
2
0
1965
1966
1967
1968
149
150
25
Energy flows through ecosystems
Different types of producers
–
–
Photo-autotrophs –
CO2+ H2O Æ C6H12O6
(Energy provided by solar energy)
Chemo-autotrophs–
CO2 or CH4 Æ Æ Æ C(H2O)n
(Energy provided by H2S+ O2 Æ
Energy)
151
Trophic level refers to
the feeding level of an organism
Different types of consumers
–
–
–
152
Primary consumers – herbivores,
grainivores, frugivores,
“Higher” consumers – carnivores,
insectivores, parasites,
Detritivores -- break down dead organic
material
ƒ Scavengers – Vultures and crabs
ƒ Decomposers – Bacteria and fungi
153
154
Primary Productivity
•
•
Green plants capture about 1% of the
solar energy that falls on their leaves.
In a given area during a given period of
time …
– Gross primary productivity (GPP) is the
total organic matter produced by
producers.
– Net primary productivity (NPP) is the
amount of organic matter produced that
is available to consumers.
– Biomass is the total mass of all
organisms.
Primary
productivity
of various
ecosystems.
155
156
26
Primary productivity of various ecosystems.
Low
Exercise:
Why do they
have different
NPP ?
High
157
158
Primary Productivity
We did not talk about the material covered in
the following 20 slides.
But they will show up in the final examine.
•
•
•
The productivity in a given area is ultimately
determined by the amount of energy it receives.
Thus, NPP often increases as the growing season
lengthens.
In aquatic ecosystems, light and nutrients limit
primary production.
In terrestrial ecosystems, temperature, moisture,
and nutrients limit primary production.
159
160
On a local scale, nutrients in the soil can
play key roles in limiting primary production.
Not all NPP is consumed
•
•
•
161
NPP not consumed will eventually become
available to decomposers.
And, not 100% of the NPP consumed by
herbivores is assimilated
Herbivores assimilate about 20% of
consumed biomass; carnivores 5%. On
average, 10% of consumed biomass is
assimilate by consumers.
162
27
~10% assimilation
from one level to the next
The assimilation by consumers is only ~10%
17%
Growth
= assimilation
33%
Cellular
respiration
50%
Feces
= heat loss
163
Exercise: Why most food chains contain
less than 4 trophic levels?
Primary
producer
Primary
consumer
Secondary
consumer
164
Diminishing energy due to heat loss
Primary
producer
Tertiary
consumer
Primary
consumer
Secondary
consumer
Tertiary
consumer
One-way energy flow
165
166
Energy availability (NPP)
affects
the length of a food chain.
Diminishing energy
due to heat loss
Fig. 54.1
167
168
28
Ecological Pyramids
Exercise:
Why are top
predators rarer
than lower
consumers?
Moving up a food chain, you generally find
fewer individuals, lower total biomass, and
lower total energy at each successive
trophic level.
169
Pyramid of numbers
1
11
170
Pyramid of biomass
Decomposer
(5 g/m2)
Carnivore
Herbivore
Second-level carnivore
(1.5 g/m2)
First-level carnivore (11 g/m2)
4,000,000,000
Herbivore (37 g/m2)
Photosynthetic plankton
Photosynthetic plankton (807 g/m2)
171
Here is a good reason to be a vegetarian
Pyramid of energy
Decomposer
(3890 kcal/m2/yr)
172
First-level carnivore
(48 kcal/m2/yr)
Herbivore
(596 kcal/m2/yr)
Photosynthetic plankton
(36,380 kcal/m2/yr)
173
174
29
Trophic cascade
Three-level top-down cascade
Trout – Invertebrate -- Algae
•
The effect of one trophic level flows
up or down to other levels.
Top-down effects
– Bottom-up effects
–
175
No Trout
No Trout
176
Algae-eating insect
Algae (g/m2)
Algae
Exercise:
Fish added
No fish added
In a imaginary state, people love to hunt –
deer hunting. They hate the wolves that
take “their” deer. Wolves are bad. So they
do everything to get rid of wolves.
200
100
0
60
50
40
30
20
10
0
5000
4000
3000
2000
1000
0
What would happen if wolves disappeared?
before
after
fish addition
177
Sand County Almanac
178
Bottom-up effects
Three levels
By Aldo Leopold
I have lived to see state after state extirpate its wolves.
I have watched the face of many wolfless mountain,
and seen the south-facing slopes wrinkle with a maze of
new deer trails.
I have seen every desuetude, and then to death.
I have seen every edible tree defoliated to the height of a
saddle horn.
Increase the amount of light,
increase predator biomass
179
Predator biomass
Damselfly nymph
300
Vegetation biomass Herbivore biomass
Predatory fish
Chironomids
Damselflies
(number/g algae) (number/m2)
Four-level top-down cascade
Amount of light
180
30
Bottom-up effects
Four levels
Exercise: Why are there
portions of the curves
where vegetation
biomass does not
increase as productivity
increases?
Increase productivity,
increase food-chain
length
Productivity
181
Productivity
182
31