Variation in species interaction outcomes
Scott Chamberlain
28 September 2012
Shortcuts:
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Bronstein 1994 TREE
Interaction outcomes are important for:
Individual growth and reproduction - e.g., herbivory
Population growth - e.g., fox and hare
Formation and dynamics of food webs - e.g., interaction strengths important for stability
Evolution - e.g., interactions alter traits important for interaction
Mean interaction outcomes
What about variation in outcomes?
Within a species interaction, outcomes are not just +, 0, or -
Outcomes within a species interaction can range from - to +
What is meant by variation in interaction outcomes?
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Not error variation in outcome in one context
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The variation in outcomes among more than one context
Variation in interaction outcomes - example
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Not error variation in outcome in one context
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The variation in outcomes among more than one context
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And so on for other types of contexts: time, space, etc.
Images from Todd Palmer
Two points:
- Variation in outcomes is common
- Variation in outcomes is important
Variation in outcomes is common
Mutualism
Herbivory outcome for Acacia trees varies with ant species identity (Palmer et al. 2008)
Variation in outcomes is common
Competition/facilitation
Outcomes vary from competition at low elevation to facilitation at high elevation (Callaway et al. 2002)
And people are calling for examination of variation
To the extent that different biotic interactions are important contributors to different population dynamics within a species, variable interactions might be expected to be a powerful such source. - Joseph Travis 1996 AmNat
The full range of natural outcomes...may reveal far more about...ecological and evolutionary dynamics than does the average outcome. - Judith Bronstein 1994 TREE
...understanding context-dependency is critical for such reconciliation [of variation in species interaction outcomes]. - Anurag Agrawal et al. 2007 FREE
Questions
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How do different species interaction types differ in variation in outcomes?
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What are the relative importance of drivers of variation in outcomes?
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Does variation in outcome differ by ecosystem type?
Two major distinctions can be made:
- Types of species interactions, and
- Gradients along which interaction occur
How do different species interaction types differ in variation in outcomes?
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Mean strength
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Mutualisms weaker than antagonisms (Morris et al. 2007)
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General sense in literature that mutualisms less important because so variable (Sachs & Simms 2006)
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Weak interactions the most variable (Berlow et al. 1999)
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Interaction complexity
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Predation more specialized than mutualism (Gomez et al. 2010)
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Strength greater with fewer interactions (Edwards et al. 2010)
Gradients that drive variation in interaction outcome
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Same two species
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Abiotic: Nutrients
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Space: Across sites
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Time: Across hours, days, or years
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Third party presence: Two species w/ or w/o a third species
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Just one species is the same
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Species identity: Sp. A interacts with either sp. B or sp. C
Meta-analysis
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Web of Science search
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Experimental studies only
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Interaction outcome w/ & w/o competitor, predator, or mutualist
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Error estimates & sample sizes available
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Response variables: abundance, population growth, reproduction, etc.
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Responses measured over >1 year, population, or species, etc.
E.g. Getting data from a single study..cont.
Data analyses
Weighted general linear models
Magnitude (CV)
CV ~ context type + species interaction type + err
CV ~ context type + species interaction type + ecosystem + err
Sign change
sign change ~ context type + species interaction type + err
sign change ~ context type + species interaction type + ecosystem + err
Final dataset - 353 papers
How do different species interaction types differ in variation in outcomes?
How do different species interaction types differ in variation in outcomes?
How do different species interaction types differ in variation in outcomes?
Why is mutualism more variable?
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I don't know, but...
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A few possibilities:
Conditionality seems to be discussed more in the mutualism literature, but perhaps just what mutualism people are interested in?
Possible that species interactions are misidentified? -> Predation is likely predation, whereas not so for mutualism and competition
Mutualism is just more variable = but what is it that makes this so? Any ideas?
What are the relative importance of drivers of variation in outcomes?
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Same two species
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Abiotic: Nutrients
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Space: Across sites
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Time: Across hours, days, or years
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Third party presence: Two species w/ or w/o a third species
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Just one species is the same
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Species identity: Sp. A interacts with either sp. B or sp. C
What are the relative importance of drivers of variation in outcomes?
** No difference for sign change
What does this mean?
Species identity > third party presence
Perhaps it's really species identity that matters when considering community context
Spatial tended to be > temporal variation, but not significant
Interesting that just as much variation in interactions can occur in space as in time
Drivers of variation in outcomes X interaction type? - Sign
Drivers of variation in outcomes X interaction type? - Sign
Mutualism > predation & competition along spatial and 3rd party axes
Makes sense at least in some mutualisms, e.g. ->
Drivers of variation in outcomes X interaction type? - Sign
Predation < competition & mutualism along abiotic and temporal axes
Predator-prey with largely mobile species, escape abiotic constraints
Mobility
Results qualitatively the same when accounting for mobility
Reasoning for ecosystem differences
Oceans buffer abiotic variables more so than freshwater and marine
Which could lead to less variation in interaction outcomes in oceans
Any ideas?
Reid et al. 2009
Conclusions
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Types of species interactions differed in outcome variation
We can't treat different species interactions as equivalent
In interaction webs, it may be most important to understand variation in mutualistic links
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Types of gradients differed in outcome variation
Some sources of variation in species interactions should be given priority (i.e., species identity), especially in new study systems
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Types of ecosystems differed in outcome variation
Outcome variation: terrestrial >= freshwater > marine
Future work
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Add other species interaction types
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Do any variables correlate with variation in species interaction outcomes?
Do body size ratios predict variable outcomes?
How does variation in outcome vary with latitude?
Interaction outcomes are thought to increase in strength nearer the equator (Schemske et al. 2009)
Stronger biotic interactions nearer equator => Greater biodiversity
What about variation?
Stronger mean interactions are thought to be associated with less variation (Berlow 1999)
Stronger mean nearer equator => less variable nearer equator
How does variation in outcome vary with latitude? - Predation
Latitude
Variation may depend on latitude, or some variable correlated with latitude
Variation in outcomes may increase in some areas more than others with climate change
What about latitude is correlated with variation in interaction outcomes?
Exploring this question with Jeremy Kerr (Univ. Ottawa), looking at:
Temperature
Precipitation
Etc.
How can we test these findings in the field?
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What are the consequences of variation in outcomes?
Probably very difficult to manipulate variation in outcomes
Modeling perhaps?
Any ideas?
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What are relative importance of mechanisms for variation in outcome?
A bit easier
Manipulate many gradients discussed here:
Abiotic gradients (e.g., nutrients)
Third party presence (e.g., presence of predator in plant-herbivore interaction)
Thanks to...
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Data
Tens of authors who provided data
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Data collection
Yosuke Akiyama, Neha Deshpande, Rohini Sigireddi, Prudence Sun, Edward Realzola, Rameez Anwar, Matt King, Roy Wu, Than Vu
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Discussion
The Rudgers-Whitney lab, Jennifer Rudgers, Ken Whitney, Volker Rudolf, Dennis Cox, Steve Hovick, Tom E.X. Miller, Elizabeth Elle, Lisandro Benedetti-Cecchi
Taxonomic bias
Though can't account for it due to lack of replication across levels
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