Biodiversity and Ecosystem

Function
 What is biodiversity?
• OED: “biodiversity Ecol., diversity of plant and
animal life, as represented by the number of
extant species”
• Ricklefs & Miller: Biodiversity includes a
number of different levels of variation in the
natural world: genetic, species, ecosystem
• Begon et al. “The term may be used to describe
the number of species, the amount of genetic
variation or the number of community types
present in an area”.
• But…most studies do focus on species diversity
Are Aspects of Ecosystem
Functioning Dependent on
Biodiversity?
 Functional Consequences of
Biodiversity:

Numbers and Organismal Ecosystem

Kinds of traits Processes
Species
 Traits and Ecosystem
Function
• Traits may mediate energy and material
flow directly
• Traits may alter abiotic conditions (limiting
resources, disturbance, microclimate)
 Trait Expression Is
Determined By:
• Species richness
• Species evenness
• Species composition
• Species interaction
• Temporal and spatial variation
Photo: Arctic LTER
 Leaf area index
2.0 Pteridophyte
Evergreen
LAI, one-sided (m m )
-2

Deciduous
1.5 Graminoid
2

1.0

0.5

0.0
5 1 2 H 94-4 3 4 6 HV 13 14 15 16 9
Barrens Wetland Heath MNT Moist acidic tundra Shrubs
 2
r =0.93
3.2

Total foliar N (g m ground area)

2.8

2.4

2.0
-2

1.6

1.2

0.8

0.4

0.0
0.0 0.4 0.8 1.2 1.6 2.0 2.4
LAI
 The Ecosystem/Ecology
Divide
• Key ecosystem types in Arctic tundra show clear
differences in key species and functional types
• But at the ecosystem level there are clear patterns
in the landscape irrespective of species
composition
• Bulk measures like LAI and foliar N are good
descriptors of process rates
• So, are species important?
 An exercise in pairs
• Identify specific examples of links between
biodiversity and ecosystem function/process
 Species Richness and
Ecosystem Function:
Theory

• If niches are complementary, adding species

could increase process rates linearly
• As niches overlap the response should
saturate
Niche differentiation and productivity. a,
A simple model — the 'snowballs on the
barn' model — of niche differentiation
and coexistence. The range of conditions
in which each species can exist is shown
with a circle, the position of which is
defined by its centre. By randomly
choosing locations for various numbers
of circles (species), it is possible to
calculate the effect of diversity on the
'coverage' of the heterogeneous habitat.
The amount of such coverage is
proportional to community biomass. b,
Results of simulations (triangles) and of
an analytical solution (solid curve) to the
effects of diversity on community
productivity for the snowballs on the
barn model

From: Tilman (2000), Nature.

Tilman’s biodiversity experiment
(A) Dependence of
1996 aboveground plant biomass
(that is, productivity) (mean and
SE) on the number of plant
species seeded into the 289 plots.

(B) Dependence of 1996 above-

ground plant biomass on the
number of functional groups
seeded into each plot. Curves
shown are simple asymptotic
functions fitted to treatment
means. More complex curves did
not provide significantly better
fits

From: Tilman et al. (1997) Science

 Problems With Richness
Experiments
• Disentangling interactions in natural systems is
difficult
• Measuring productivity (below ground)
• Scale (too short and too small)
• ‘Sampling effect’ problem in constructed
communities
– Sampling effect may be an important biological
property or an experimental artefact if natural
community assembly rules are broken
Hypothesized mechanisms
involved in biodiversity
experiments using synthetic
communities. Sampling effects
are involved in community
assembly, such that communities
that have more species have a
greater probability of containing a
higher phenotypic trait diversity.
Phenotypic diversity then maps
onto ecosystem processes through
two main mechanisms:
dominance of species with
particular traits, and
complementarity among species
with different traits. Intermediate
scenarios involve
complementarity among
particular species or functional
groups or, equivalently,
dominance of particular subsets
of complementary species.

From: Loreau et al (2001)

Science
 Richness Conclusions
• With our knowledge now, we cannot reject
the hypothesis that a few dominant species
suffice to provide the functional diversity
that is necessary to explain the level of
primary production observed in grassland
ecosystems at the small spatial and temporal
scales considered in recent experiments.
 Species Evenness
• Human effects on species more commonly
involve alteration of relative abundance
than extinction
• Little research on importance of evenness of
function so far
• Future richness experiments should include
evenness effects
 Species Composition:
• Species mediate pathways of energy
and material flow
• Examples: Introduced species can alter
patterns of ecosystem processes
 Introduced Species Can Alter
Patterns of Ecosystem Processes I

• Introduction of N-fixing tree Myrica faya to

N-limited Hawaiian forests led to 5-fold
increase in N inputs
• Significant impacts on forest structure and
function

Vitousek et al. (1987) Science

 Introduced Species Can Alter
Patterns of Ecosystem Processes II
• Introduction of deep-rooted salt cedar
(Tamarix sp.) to Mojave and Sonaran
deserts resulted in:
– Increased water accessed by vegetation
– Increased surface litter and salts
– Inhibited many native species, reduced
biodiversity

Berry (1970)
 Introduced Species Can Alter
Patterns of Ecosystem Processes III

• Introduction of Agropyron cristatum,

tussock grass, to US Great Plains
– Reduced allocation to roots compared to
native grasses
– Soil N levels reduced, and 25% less total
soil C compared to native prairie soil

•Christian & Wilson (1999)

 Introduced Species Can Alter
Patterns of Ecosystem Processes IV
• Introduction of Bromus tectorum,
cheatgrass, to western US
– Fire frequency increased by a factor of 10 in
the >40 million ha it now dominates

•Whisenant (1990)
 CASE STUDY:
Diversity of tropical trees
and carbon water relations
 Exploring compositional effects on
ecosystem function

• Are Amazonian trees

water stressed?
• Does water stress
depend on soil texture?
• Does water stress
depend on species?
Ecological
Diversity in
Amazonian
Rain Forest
Diversity in Rooting Depths
Dawn water potential along
2.0
a soil texture gradient
x -1)

1.5
Dawn leaf water potential (MPa

1.0

0.5

0.0

-0.5
3 1 4 5 6 9 10 2 11 12 7 8 13
High clay High sand
Site
Williams et al (2002)
Species Interactions
• Mutualism
• Trophic interaction
– Predation
– Parasitism
– Herbivory
• Competition
 Mutualism
• N-fixation in plant-microbe symbiosis
• Plant-mycorrhizal associations
– Both increase production and accelerate
succession
• Decomposition is driven by highly
integrated consortia of microbes
 CASE STUDY
Mycorrhizal fungal diversity
determines plant biodiversity,
ecosystem variability and
productivity
 Background
• arbuscular mycorrhizal fungi (AMF) form
mutualistic symbiotic associations with
the roots of 80% of all terrestrial plant
species, thereby acting as extensions of
plant root systems and increasing
nutrient uptake, especially of
phosphorus
• Communities vary in AMF biodiversity
 Experimental design
• Expt 1: Greenhouse experiment, 48
microcosms simulating European
calcareous grassland
– Quantified plant growth with no AMF or
different diversities of AMF
• Expt 2: Field experiment, 70
macrocosms simulating N. American old
fields.
– 15 plant species per plot, random mix of
different numbers of species (out of 23)
AMF, quantified primary productivity
Experiment 1 van der Heijden 1998
Experiment 2 van der Heijden 1998
 More diverse microbial
communities are more efficient
Soil microbial functional diversity
(Shannon index H') and metabolic
quotient (qCO2 = soil basal
respiration/soil microbial biomass)
correlate inversely. A higher
diversity in the organic plots is
related to a lower qCO2, indicating
greater energy efficiency of the more
diverse microbial community. The
Shannon index is significantly
different between both conventional
systems (CONFYM, CONMIN) and
the BIODYN system, the qCO2,
between CONMIN and BIODYN
(P < 0.05).
Maeder 2002
 Trophic Interactions
• Modify fluxes of energy and materials
• Influence abundance of species that control
these fluxes
– e.g., predator removal can lead to a cascade of
ecological effects
(A) Changes in sea otter abundance
over time at several islands in the
Aleutian archipelago and concurrent
changes in (B) sea urchin biomass,
(C) grazing intensity, and (D) kelp
density measured from kelp forests at
Adak Island. Error bars in (B) and (C)
indicate 1 SE. The proposed
mechanisms of change are portrayed
in the marginal cartoons--the one on
the left shows how the kelp forest
ecosystem was organized before the
sea otter's decline and the one on the
right shows how this ecosystem
changed with the addition of killer
whales as an apex predator. Heavy
arrows represent strong trophic
interactions; light arrows represent
weak interactions.

Estes et al. (1998) Science

Experimental Lakes
•Two lakes dominated by
Zooplanktivorous
fishes(minnows)
•And two other lakes
dominated by Piscivorous
fish (bass)
•Of each pair, one was
fertilized with N+P while
the other was left as a
control

Schindler et al (1997) Science

bass + P  minnows, grazers, algae,
lake shifts from C source to sink
Comparison of crustacean
grazer length, primary
production, and DPCO2
[PCO2(lake) - PCO2(air)] before,
during, and after a period of low
minnow abundance in Peter
Lake (1994 to 1995). Data are
shown as the mean ± SD.
Sample sizes are given in
parentheses. Dotted line in
bottom panel represents
dissolved CO2 in equilibrium
with the atmosphere
Schindler et al (1997) Science
 Trophic Interactions:
Conclusions
• All types of organisms must be considered
in understanding biodiversity effects
• Interactions among species must be
considered
• Changes in interactions can alter traits
expressed by species, so presence/absence
of species is insufficient to predict impact
 Biodiversity and Ecosystem
Services
• Ecosystem services are defined as the processes
and conditions of natural ecosystems that support
human activity and sustain human life
• E.g., maintenance of soil fertility, climate
regulation, natural pest control
• E.g., flows of ecosystem goods such as food,
timber and freshwater
Attaching Value to Biodiversity
• Techniques used include direct valuation
based on market prices, and estimates of
what individuals are willing to pay to
protect endangered wildlife
• Valuation of marginal losses that
accompany specific biodiversity changes
are most relevant to policy decisions
• Predictions are highly uncertain
What you should have learned
today
• The ways in which biodiversity can affect
ecosystem function
• Experimental approaches and the sampling
effect
• Examples of species traits that control
particular processes
• The concept of ecosystem services and
valuation
 References
• Chapin et al (2000) Consequences of changing
biodiversity. Nature 405: 234-242
• Tilman, Wedin and Knops (1996) Productivity and
sustainability influenced by biodiversity in grassland
ecosystems. Nature 379: 718-720
• Naeem & Li (1997) Biodiversity enhances ecosystem
reliability. Nature 390:507-509
• Van der Heijden et al. (1998) Nature 396: 69-72
• Constanza et al (1997) The value of the world's
ecosystem services and natural capital. Nature 387:
253-260
• Maeder et al (2002) Soil fertility and biodiversity in
organic farming. Science 296: 1694-7
 Reading for next week
• Pfisterer. A.B. & B. Schmid. 2002. Diversity-dependent
production can decrease the stability of ecosystem
functioning. Nature 416 84-86
– what insights does this experiment provide?
– what are the criticisms of the approach?
• McCann, K., A. Hastings G. R. Huxel. 1998. Weak trophic
interactions and the balance of nature. Nature 395 794-8
– what insights does the modelling provide?
– what are the criticisms of the approach?