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Cascading effects of spatial structure
across trophic levels
Rachel M. Germain
Killam/Biodiversity Postdoctoral Fellow
Main collaborators: Natalie Jones, Tess Grainger, Ben Gilbert, Andrew MacDougall
Community ecologist: species coexistence
and metacommunities
Schematic modified from Velland 2016
Botany seminar Oct. 11th @12:30pm
env gradient
Collaborators
Tess Grainger Natalie Jones Ben Gilbert
Biological communities rarely exist in isolation
‘metacommunity’
biotic/abiotic environment
Predictions of island biogeography/metapopulation
theories
species richness/
site occupancy
larger, less isolated patches
species richness/
site occupancy
smaller, more isolated patches
MacArthur & Wilson 1967; Hanski 1994
Predictions of island biogeography/metapopulation
theories
species richness/
site occupancy
larger, less isolated patches
species richness/
site occupancy
smaller, more isolated patches
MacArthur & Wilson 1967; Hanski 1994
Spatial constraints likely vary among species
sessile vs. motile, dispersal ability, cognitive function → behavior (habitat selection, territory, predator
avoidance), body size, matrix sensitivity, etc.
P
C2
C1
C3
Cascading spatial dynamics among trophic
levels patch
isolation
patch
size
spatial patterns
spatial patterns
Multi-trophic extensions of island biogeography
and metacommunity theories
Trophic dependency: predators can only establish in patches that
already contain their prey
• prey must outpace their predators
• predator distributions should be a subset of prey distributions
• IBT patterns should be stronger at higher trophic levels
Holt 2002 EcoRes; Leibold et al. 2004 EcoLett; Harvey & MacDougall 2014 Ecology
Assumes that predators: are specialists, are bound to same habitat patches as their prey, and experience
the world at the same spatial scales as their prey
What are the consequences for diversity in systems where
spatial constraints differ within and among trophic levels?
aspen-understory plants
Jones, Germain, et al. 2015 JEcology
tall-grass prairie plants
Germain et al. 2013 AmNat
milkweed insect specialists
Grainger, Germain, et al.
in review Ecology
What are the consequences for diversity in systems where
spatial constraints differ within and among trophic levels?
aspen-understory plants
Jones, Germain, et al. 2015 JEcology
tall-grass prairie plants
Germain et al. 2013 AmNat
milkweed insect specialists
Grainger, Germain, et al.
in review Ecology
Tall-grass prairie restoration efforts in SW Ontario
System: tall-grass prairie plant communities
System: tall-grass prairie plant communities
How does risk avoidance
behavior by small mammals
affect spatial patterns in plant
communities?
predation
risk
cognition
yes
yes
no
range
size
Predation risk increases with distance from
old-field edge
pre-
prairie
more removal less removal
old-
field
Prediction:
Closed
old-field
Pre-prairie
-30 -10 10 30 50 70 90+
Distance from edge (m)
Prop.seedsremoved
F1,84 = 150.9, P < 0.0001*
Seed removal was strongest in the old-field, and
decreased with distance from the edge
Consequences of small mammals for plant
diversity depends on foraging selectivity
Cafeteria trial w/ 8 plant species from 3
functional groups
B. kalmii H. divaricatus R. blanda
0
2
4
6
8
10Seedsremaining
Seed species
Forb seeds were consumed significantly more than shrub and
grass seeds
Functional group: F2,23 = 29.97, P < 0.001*
Species: F5,23 = 0.35, P = 0.876
Forbs Shrubs Grasses
P = 0.012*F1,48 = 12.82, P <0.001*
β-diversity(Jaccard’sdissimilarity)
α-diversity(speciesperplot)
α β
Distance from edge (m) Distance from edge (m)
What does this all mean for plant community
distributions?
What does this all mean for plant community
distributions?
P = 0.012*F1,48 = 12.82, P <0.001*
β-diversity(Jaccard’sdissimilarity)
α-diversity(speciesperplot)
α β
Distance from edge (m) Distance from edge (m)
oldfield
increasing
β-diversity
decreasing
seed
predation
prairie interior
Net effect: emergent patterns in plant diversity
that reflect risk environment
What are the consequences for diversity in systems where
spatial constraints differ within and among trophic levels?
aspen-understory plants
Jones, Germain, et al. 2015 JEcology
tall-grass prairie plants
Germain et al. 2013 AmNat
milkweed insect specialists
Grainger, Germain, et al.
in review Ecology
System: milkweed insect specialists
habitat use
generalist
milkweed
specialist
Milkweed specialists differ in susceptibility to
predators and dispersal ability
Constrained
by predation1:
yes
1Duffey & Scutter 1972, Zalucki & Kitching 1982, Smith et al. 2008
2Jones & Parella 1986, St Pierre & Hendrix 2003, McCauley et al. 1981
no
yes no
Constrained
by size/isolation2:
<100 m >1000 m
Predictive framework for island biogeography
informed by natural history
P, S, I
S, Inone
P
Patchoccupancy
Patch size/isolation
P = predator, S = size, I = isolation
Do constraints of patch
size/isolation vary among
species, and are they altered
by predation?
Field surveys of milkweed patches
Patch isolation
Patch size (m2)
𝐼𝑖 = 1 −
𝑗≠𝑖
𝑛
𝐴𝑗 𝑒
−𝑑 𝑖𝑗 𝛼
Hanski 1994 TREE
Presence/absence data Predator abundances
occupancy ~ patch size x patch isolation x predator abundances
Predictive framework for island biogeography
informed by natural history
none
P
N.S.
Patch isolation
Occupancy
P, S, I
S, I
P = predator, S = size, I = isolation
Predictive framework for island biogeography
informed by natural history
none
P
P = predator, S = size, I = isolation
P, S, I
S, I
Predictive framework for island biogeography
informed by natural history
none
P
P = predator, S = size, I = isolation
P, S, I
S, I
Zalucki & Kitching 1982a JZoology, Zalucki & Kitching 1982b Oecologia
“More eggs were laid per plant on single
isolated plants than on plants within a patch”
“The trend was for increasing [mortality] with
increasing patch size” → predation
Predictive framework for island biogeography
informed by natural history
none
P
P = predator, S = size, I = isolation
P, S, I
S, I
Tying it all together…
Dispersal-limited species were directly
affected by spatial constraints
Predation affected palatable species
indirectly through interactions with
spatial drivers
Incorporating predator-avoidance
tactics (refuge seeking and crypsis) can
help refine predictions
Biogeographic
constraints highly
variable among species
in this community – but
fairly predictable
Matrix-dwelling predators
can weaken or modify the
effects of spatial drivers
What are the consequences for diversity in systems where
spatial constraints differ within and among trophic levels?
aspen-understory plants
Jones, Germain, et al. 2015 JEcology
tall-grass prairie plants
Germain et al. 2013 AmNat
milkweed insect specialists
Grainger, Germain, et al.
in review Ecology
System: Aspen understory plant communities
Do effects of stand size and isolation on species
distributions vary with dispersal mode?
cognition
yes
no
range
size
Field surveys of aspen stands
𝐼𝑖 = 1 −
𝑗≠𝑖
𝑛
𝐴𝑗 𝑒
−𝑑 𝑖𝑗 𝛼
Hanski 1994 TREE
Stand characteristics
size (m2)
isolation
stand age
Presence/absence data Dispersal mode
No aid
Wind
Animal
Aspen stand
Adjacent grassland
Predicted effects of stand size and isolation
for different dispersal modes
No aid Wind-dispersed Animal-dispersed
Observed effects of stand size and isolation
for different dispersal modes
No aid Wind-dispersed Animal-dispersed
competition among dispersal modes?
ruderal species and disturbance?
Paired plot design to identify aspen specialists
vs. generalists or grassland specialists
Quantifying species affinity for aspen stands
all species included
occurs in aspen stands ≥50% of the time
occurs in aspen stands ≥ 66% of the time
occurs in aspen stands ≥75% of the time
cut-off is not always clear in “diffuse” metacommunities
Cook et al. 2002 EcoLett; Leibold et al. 2004 EcoLett
metacommunity = species that occur in favourable habitat
patches imbedded in a matrix of unfavourable habitat
The results are insensitive to cut-off, except
when all species are included
SR ~ stand size
Matrix-associated species can obscure metacommunity processes – paired plot design is one
day to discount their effects
IBT patterns are somewhat consistent with
our initial predictions, but …
There are three complicating factors:
• dispersal mode
• unexplained inconsistencies
• grassland specialists + generalists
…and so, to summarize…
ability to persist in matrix habitat = modifications to biogeographic
constraints in insect species that are sensitive to predators
greater habitat use by granivores in low risk areas = gradients in
ecological filters that dictate plant species distributions
habitat selection by animals = non-IBT patterns for animal-dispersed
plant species
“patch area and isolation are surprisingly poor
predictors of occupancy for most species”
data from 1,015 bird, mammal, reptile, amphibian, and
invertebrate population networks
metapopulation metacommunity∑ =
Island biogeography in non-island systems
suitable habitat unsuitable habitat
Island biogeography in non-island systems
suitable habitat unsuitable habitat
Are patch size and isolation unimportant compared to/in combination with local factors? How do spatial
dynamics cascade across trophic levels? How do spatial dynamics change with species additions or losses at
different trophic levels? Should conservationists only be focusing on patch size/isolation?
Acknowledgements
Co-authors
Natalie Jones
Tess Grainger
Ben Gilbert
Aaron Hall
Lynn Baldwin
Andrew MacDougall
Karl Cottenie
Laura Johnson
Elizabeth Gillis
Webpage: rgermain.wordpress.com
Email: rgermain@zoology.ubc.ca
Office: BIODIV245
Monarch caterpillar mortality as a function of
stand size
PCoA testing turnover in proportion of
species belong to each dispersal modes
UBC BLISS talk 2016

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UBC BLISS talk 2016

  • 1. Cascading effects of spatial structure across trophic levels Rachel M. Germain Killam/Biodiversity Postdoctoral Fellow Main collaborators: Natalie Jones, Tess Grainger, Ben Gilbert, Andrew MacDougall
  • 2. Community ecologist: species coexistence and metacommunities Schematic modified from Velland 2016 Botany seminar Oct. 11th @12:30pm env gradient
  • 4. Biological communities rarely exist in isolation ‘metacommunity’ biotic/abiotic environment
  • 5. Predictions of island biogeography/metapopulation theories species richness/ site occupancy larger, less isolated patches species richness/ site occupancy smaller, more isolated patches MacArthur & Wilson 1967; Hanski 1994
  • 6. Predictions of island biogeography/metapopulation theories species richness/ site occupancy larger, less isolated patches species richness/ site occupancy smaller, more isolated patches MacArthur & Wilson 1967; Hanski 1994
  • 7. Spatial constraints likely vary among species sessile vs. motile, dispersal ability, cognitive function → behavior (habitat selection, territory, predator avoidance), body size, matrix sensitivity, etc.
  • 8. P C2 C1 C3 Cascading spatial dynamics among trophic levels patch isolation patch size spatial patterns spatial patterns
  • 9. Multi-trophic extensions of island biogeography and metacommunity theories Trophic dependency: predators can only establish in patches that already contain their prey • prey must outpace their predators • predator distributions should be a subset of prey distributions • IBT patterns should be stronger at higher trophic levels Holt 2002 EcoRes; Leibold et al. 2004 EcoLett; Harvey & MacDougall 2014 Ecology Assumes that predators: are specialists, are bound to same habitat patches as their prey, and experience the world at the same spatial scales as their prey
  • 10. What are the consequences for diversity in systems where spatial constraints differ within and among trophic levels? aspen-understory plants Jones, Germain, et al. 2015 JEcology tall-grass prairie plants Germain et al. 2013 AmNat milkweed insect specialists Grainger, Germain, et al. in review Ecology
  • 11. What are the consequences for diversity in systems where spatial constraints differ within and among trophic levels? aspen-understory plants Jones, Germain, et al. 2015 JEcology tall-grass prairie plants Germain et al. 2013 AmNat milkweed insect specialists Grainger, Germain, et al. in review Ecology
  • 12. Tall-grass prairie restoration efforts in SW Ontario
  • 13. System: tall-grass prairie plant communities
  • 14. System: tall-grass prairie plant communities How does risk avoidance behavior by small mammals affect spatial patterns in plant communities? predation risk cognition yes yes no range size
  • 15. Predation risk increases with distance from old-field edge pre- prairie more removal less removal old- field Prediction:
  • 16. Closed old-field Pre-prairie -30 -10 10 30 50 70 90+ Distance from edge (m) Prop.seedsremoved F1,84 = 150.9, P < 0.0001* Seed removal was strongest in the old-field, and decreased with distance from the edge
  • 17. Consequences of small mammals for plant diversity depends on foraging selectivity Cafeteria trial w/ 8 plant species from 3 functional groups B. kalmii H. divaricatus R. blanda
  • 18. 0 2 4 6 8 10Seedsremaining Seed species Forb seeds were consumed significantly more than shrub and grass seeds Functional group: F2,23 = 29.97, P < 0.001* Species: F5,23 = 0.35, P = 0.876 Forbs Shrubs Grasses
  • 19. P = 0.012*F1,48 = 12.82, P <0.001* β-diversity(Jaccard’sdissimilarity) α-diversity(speciesperplot) α β Distance from edge (m) Distance from edge (m) What does this all mean for plant community distributions?
  • 20. What does this all mean for plant community distributions? P = 0.012*F1,48 = 12.82, P <0.001* β-diversity(Jaccard’sdissimilarity) α-diversity(speciesperplot) α β Distance from edge (m) Distance from edge (m)
  • 21. oldfield increasing β-diversity decreasing seed predation prairie interior Net effect: emergent patterns in plant diversity that reflect risk environment
  • 22. What are the consequences for diversity in systems where spatial constraints differ within and among trophic levels? aspen-understory plants Jones, Germain, et al. 2015 JEcology tall-grass prairie plants Germain et al. 2013 AmNat milkweed insect specialists Grainger, Germain, et al. in review Ecology
  • 23.
  • 24. System: milkweed insect specialists habitat use generalist milkweed specialist
  • 25. Milkweed specialists differ in susceptibility to predators and dispersal ability Constrained by predation1: yes 1Duffey & Scutter 1972, Zalucki & Kitching 1982, Smith et al. 2008 2Jones & Parella 1986, St Pierre & Hendrix 2003, McCauley et al. 1981 no yes no Constrained by size/isolation2: <100 m >1000 m
  • 26. Predictive framework for island biogeography informed by natural history P, S, I S, Inone P Patchoccupancy Patch size/isolation P = predator, S = size, I = isolation Do constraints of patch size/isolation vary among species, and are they altered by predation?
  • 27. Field surveys of milkweed patches Patch isolation Patch size (m2) 𝐼𝑖 = 1 − 𝑗≠𝑖 𝑛 𝐴𝑗 𝑒 −𝑑 𝑖𝑗 𝛼 Hanski 1994 TREE Presence/absence data Predator abundances occupancy ~ patch size x patch isolation x predator abundances
  • 28. Predictive framework for island biogeography informed by natural history none P N.S. Patch isolation Occupancy P, S, I S, I P = predator, S = size, I = isolation
  • 29. Predictive framework for island biogeography informed by natural history none P P = predator, S = size, I = isolation P, S, I S, I
  • 30. Predictive framework for island biogeography informed by natural history none P P = predator, S = size, I = isolation P, S, I S, I Zalucki & Kitching 1982a JZoology, Zalucki & Kitching 1982b Oecologia “More eggs were laid per plant on single isolated plants than on plants within a patch” “The trend was for increasing [mortality] with increasing patch size” → predation
  • 31. Predictive framework for island biogeography informed by natural history none P P = predator, S = size, I = isolation P, S, I S, I
  • 32. Tying it all together… Dispersal-limited species were directly affected by spatial constraints Predation affected palatable species indirectly through interactions with spatial drivers Incorporating predator-avoidance tactics (refuge seeking and crypsis) can help refine predictions
  • 33. Biogeographic constraints highly variable among species in this community – but fairly predictable Matrix-dwelling predators can weaken or modify the effects of spatial drivers
  • 34. What are the consequences for diversity in systems where spatial constraints differ within and among trophic levels? aspen-understory plants Jones, Germain, et al. 2015 JEcology tall-grass prairie plants Germain et al. 2013 AmNat milkweed insect specialists Grainger, Germain, et al. in review Ecology
  • 35.
  • 36. System: Aspen understory plant communities Do effects of stand size and isolation on species distributions vary with dispersal mode? cognition yes no range size
  • 37. Field surveys of aspen stands 𝐼𝑖 = 1 − 𝑗≠𝑖 𝑛 𝐴𝑗 𝑒 −𝑑 𝑖𝑗 𝛼 Hanski 1994 TREE Stand characteristics size (m2) isolation stand age Presence/absence data Dispersal mode No aid Wind Animal Aspen stand Adjacent grassland
  • 38. Predicted effects of stand size and isolation for different dispersal modes No aid Wind-dispersed Animal-dispersed
  • 39. Observed effects of stand size and isolation for different dispersal modes No aid Wind-dispersed Animal-dispersed competition among dispersal modes? ruderal species and disturbance?
  • 40. Paired plot design to identify aspen specialists vs. generalists or grassland specialists Quantifying species affinity for aspen stands all species included occurs in aspen stands ≥50% of the time occurs in aspen stands ≥ 66% of the time occurs in aspen stands ≥75% of the time cut-off is not always clear in “diffuse” metacommunities Cook et al. 2002 EcoLett; Leibold et al. 2004 EcoLett metacommunity = species that occur in favourable habitat patches imbedded in a matrix of unfavourable habitat
  • 41. The results are insensitive to cut-off, except when all species are included SR ~ stand size Matrix-associated species can obscure metacommunity processes – paired plot design is one day to discount their effects
  • 42. IBT patterns are somewhat consistent with our initial predictions, but … There are three complicating factors: • dispersal mode • unexplained inconsistencies • grassland specialists + generalists
  • 43. …and so, to summarize… ability to persist in matrix habitat = modifications to biogeographic constraints in insect species that are sensitive to predators greater habitat use by granivores in low risk areas = gradients in ecological filters that dictate plant species distributions habitat selection by animals = non-IBT patterns for animal-dispersed plant species
  • 44. “patch area and isolation are surprisingly poor predictors of occupancy for most species” data from 1,015 bird, mammal, reptile, amphibian, and invertebrate population networks metapopulation metacommunity∑ =
  • 45. Island biogeography in non-island systems suitable habitat unsuitable habitat
  • 46. Island biogeography in non-island systems suitable habitat unsuitable habitat Are patch size and isolation unimportant compared to/in combination with local factors? How do spatial dynamics cascade across trophic levels? How do spatial dynamics change with species additions or losses at different trophic levels? Should conservationists only be focusing on patch size/isolation?
  • 47. Acknowledgements Co-authors Natalie Jones Tess Grainger Ben Gilbert Aaron Hall Lynn Baldwin Andrew MacDougall Karl Cottenie Laura Johnson Elizabeth Gillis Webpage: rgermain.wordpress.com Email: rgermain@zoology.ubc.ca Office: BIODIV245
  • 48. Monarch caterpillar mortality as a function of stand size
  • 49. PCoA testing turnover in proportion of species belong to each dispersal modes