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DIVERSITY DOES NOT ALWAYS BEGET
DIVERSITY: MULTIPLE HERBIVORE
GUILDS MAINTAIN LOW WOODY
DIVERSITY IN A KENYAN SAVANNA
Grace Charles, University of California, Davis
TrumanYoung, Duncan Kimuyu, Corinna Riginos, KariVeblen
BIODIVERSITY IS DECLINING
WORLDWIDE
Plant species are also declining
UNDERSTANDING TROPHIC
INTERACTIONS AND NETWORKS CAN
HELP US UNDERSTAND DIVERSITY
PATTERNS
Scherber et al. 2010 Nature
CAN WE PREDICT HOW HERBIVORE
DIVERSITY IMPACTS WOODY DIVERSITY?
Bakker et al. 2016, PNAS
HOW DOES THE LOSS OF LARGE HERBIVORE
DIVERSITY IMPACT WOODY DIVERSITY?
• Herbivores have multiple positive and negative, direct and indirect
pathways to influence woody species diversity:
• Consumption
• Alteration of competition
• Native vs. exotic species; trees vs. grasses; dominant vs. rare species
• Habitat creation and heterogeneity
• Seed dispersal or other mutualisms
• Seed predation
• Disease spread
• Fire intensity changes
• We need to manipulate herbivore diversity not just herbivore
presence/absence
MW
C WMWC
WCO
Six combinations
of
large mammalian
herbivores
200m
C = Cattle allowed
W = Wildlife allowed
M = Mega-herbivores allowed
(elephants and giraffes)
Three
replicate
blocks
KLEE – KENYA LONG-TERM
EXCLOSURE EXPERIMENT
STUDY SITE
Semi-arid, Acacia drepanolobium
wooded grasslandKariVeblen
METHODS: HOW DOES HERBIVORE
DIVERSITY INFLUENCE WOODY
DIVERSITY?
• Mapped and recorded all woody species in each KLEE plot (6
herbivore treatments x 3 blocks)
• Allowed us to compare woody diversity in plots with 0, 1, 2, and 3
herbivore guilds
• Calculated woody species richness and Shannon diversity
• Modeled effects of herbivore treatments and herbivore
diversity (i.e., number of herbivore guilds) on woody diversity
RESULTS: HERBIVORE DIVERSITY
CAN SUPPRESS WOODY DIVERSITY
• Woody plant diversity nearly 3x higher in total exclusion plot
than plots with wildlife and megaherbivores
One"of"Jost’s"preferred"metrics"[exp(Shannon)]"
INDIVIDUAL WOODY SPECIES
RESPOND DIFFERENTLY TO
DIFFERENT HERBIVORES
0
500
1000
1500
2000
2500
3000
3500
4000
O C W WC MW MWC
Acacia drepanolobium density
0
10
20
30
40
50
60
70
80
O C W WC MW MWC
Acacia mellifera density
INDIVIDUAL WOODY SPECIES
RESPOND DIFFERENTLY TO
DIFFERENT HERBIVORES
0
10
20
30
40
50
60
70
80
90
100
O C W WC MW MWC
Grewia tenax density
INDIVIDUAL WOODY SPECIES
RESPOND DIFFERENTLY TO
DIFFERENT HERBIVORES
0
20
40
60
80
100
120
O C W WC MW MWC
Cadaba farinosa density
HERBIVORES ARE POWERFUL
SUPPRESSORS OF WOODY
DIVERSITY
Out of 20 woody species found in 2016
survey:
• 12/20 (60%) of woody species
excluded by megaherbivores (only
found in O , C, W, or WC plots)
• 5/20 (25%) of species excluded by any
wildlife herbivory (only found in O or
C plots)
• 3/20 (15%) excluded by any herbivory
(only found in O plots)
HOW IS WOODY DIVERSITY
MAINTAINED IN THIS ECOSYSTEM?
• Physical refugia
• Biological refugia
• Dispersal from
different soil type
• Important
herbivore x edaphic
interactions
• Wildlife replacement
with cattle STRI
Acacia brevispica
Black cotton soil (poorly drained) Red Soil
0
2
4
6
8
10
12
14
Acacia brevispica density
Grewia sp.
Black cotton soil (poorly drained) Red Soil
0
20
40
60
80
100
Grewia tenax density
CASCADING BIODIVERSITY LOSS:
HERBIVORE DIVERSITY SUPPRESSES
BIRD DIVERSITY
Herbivores suppress tree canopy area, in turn
reducing bird diversity
Ogada et al. 2008, Oecologia
O/C W/WC MW/MWC
A BRIEF META-ANALYSIS OF THE
LITERATURE: WHEN DOES
DIVERSITY BEGET DIVERSITY?
Criteria
• Diversity was experimentally manipulated
• 3 or more levels of diversity
• Surprisingly few studies!!
• Measured the diversity response of another group
of organisms
META-ANALYSIS: TROPHIC
RELATIONSHIPS MATTER WHEN WE
MEASURE BIODIVERSITY
• 30 tests from 20 different experiments
• Mesocosm and exclosure experiments
• 𝒳2 P = 0.01
• Intuitive result, yet rarely tested experimentally
• In cases where diversity of a higher trophic level was positively correlated
with a lower trophic level, the higher trophic level had other functions
• Seed dispersal, habitat creation
Negative	effect Positive	effect NS
Lower	trophic	level	
measured
4 2 2
Higher	trophic	level	
measured
0 13 2
CONCLUSIONS
• Herbivore diversity does not beget woody diversity
• Herbivory ≠ Herbivore diversity; lots of individual interactions
make up broader diversity patterns and relationships
• Need more studies that manipulate multiple levels of diversity
• Incorporating trophic interactions gives us a better
understanding of biodiversity patterns
KLEE:
• TrumanYoung
• KariVeblen
• Corinna Riginos
• Duncan Kimuyu
Smithsonian:
• David Kenfack
• Paul Musili
Field assistance:
• Mathew Namoni
• Jackson Ekadeli
Thanks!

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Grace Charles Ecological Society of America 2017

  • 1. DIVERSITY DOES NOT ALWAYS BEGET DIVERSITY: MULTIPLE HERBIVORE GUILDS MAINTAIN LOW WOODY DIVERSITY IN A KENYAN SAVANNA Grace Charles, University of California, Davis TrumanYoung, Duncan Kimuyu, Corinna Riginos, KariVeblen
  • 2. BIODIVERSITY IS DECLINING WORLDWIDE Plant species are also declining
  • 3. UNDERSTANDING TROPHIC INTERACTIONS AND NETWORKS CAN HELP US UNDERSTAND DIVERSITY PATTERNS Scherber et al. 2010 Nature
  • 4. CAN WE PREDICT HOW HERBIVORE DIVERSITY IMPACTS WOODY DIVERSITY? Bakker et al. 2016, PNAS
  • 5. HOW DOES THE LOSS OF LARGE HERBIVORE DIVERSITY IMPACT WOODY DIVERSITY? • Herbivores have multiple positive and negative, direct and indirect pathways to influence woody species diversity: • Consumption • Alteration of competition • Native vs. exotic species; trees vs. grasses; dominant vs. rare species • Habitat creation and heterogeneity • Seed dispersal or other mutualisms • Seed predation • Disease spread • Fire intensity changes • We need to manipulate herbivore diversity not just herbivore presence/absence
  • 6. MW C WMWC WCO Six combinations of large mammalian herbivores 200m C = Cattle allowed W = Wildlife allowed M = Mega-herbivores allowed (elephants and giraffes) Three replicate blocks KLEE – KENYA LONG-TERM EXCLOSURE EXPERIMENT
  • 7.
  • 8. STUDY SITE Semi-arid, Acacia drepanolobium wooded grasslandKariVeblen
  • 9. METHODS: HOW DOES HERBIVORE DIVERSITY INFLUENCE WOODY DIVERSITY? • Mapped and recorded all woody species in each KLEE plot (6 herbivore treatments x 3 blocks) • Allowed us to compare woody diversity in plots with 0, 1, 2, and 3 herbivore guilds • Calculated woody species richness and Shannon diversity • Modeled effects of herbivore treatments and herbivore diversity (i.e., number of herbivore guilds) on woody diversity
  • 10. RESULTS: HERBIVORE DIVERSITY CAN SUPPRESS WOODY DIVERSITY • Woody plant diversity nearly 3x higher in total exclusion plot than plots with wildlife and megaherbivores One"of"Jost’s"preferred"metrics"[exp(Shannon)]"
  • 11. INDIVIDUAL WOODY SPECIES RESPOND DIFFERENTLY TO DIFFERENT HERBIVORES 0 500 1000 1500 2000 2500 3000 3500 4000 O C W WC MW MWC Acacia drepanolobium density 0 10 20 30 40 50 60 70 80 O C W WC MW MWC Acacia mellifera density
  • 12. INDIVIDUAL WOODY SPECIES RESPOND DIFFERENTLY TO DIFFERENT HERBIVORES 0 10 20 30 40 50 60 70 80 90 100 O C W WC MW MWC Grewia tenax density
  • 13. INDIVIDUAL WOODY SPECIES RESPOND DIFFERENTLY TO DIFFERENT HERBIVORES 0 20 40 60 80 100 120 O C W WC MW MWC Cadaba farinosa density
  • 14. HERBIVORES ARE POWERFUL SUPPRESSORS OF WOODY DIVERSITY Out of 20 woody species found in 2016 survey: • 12/20 (60%) of woody species excluded by megaherbivores (only found in O , C, W, or WC plots) • 5/20 (25%) of species excluded by any wildlife herbivory (only found in O or C plots) • 3/20 (15%) excluded by any herbivory (only found in O plots)
  • 15. HOW IS WOODY DIVERSITY MAINTAINED IN THIS ECOSYSTEM? • Physical refugia • Biological refugia • Dispersal from different soil type • Important herbivore x edaphic interactions • Wildlife replacement with cattle STRI
  • 16. Acacia brevispica Black cotton soil (poorly drained) Red Soil 0 2 4 6 8 10 12 14 Acacia brevispica density
  • 17. Grewia sp. Black cotton soil (poorly drained) Red Soil 0 20 40 60 80 100 Grewia tenax density
  • 18. CASCADING BIODIVERSITY LOSS: HERBIVORE DIVERSITY SUPPRESSES BIRD DIVERSITY Herbivores suppress tree canopy area, in turn reducing bird diversity Ogada et al. 2008, Oecologia O/C W/WC MW/MWC
  • 19. A BRIEF META-ANALYSIS OF THE LITERATURE: WHEN DOES DIVERSITY BEGET DIVERSITY? Criteria • Diversity was experimentally manipulated • 3 or more levels of diversity • Surprisingly few studies!! • Measured the diversity response of another group of organisms
  • 20. META-ANALYSIS: TROPHIC RELATIONSHIPS MATTER WHEN WE MEASURE BIODIVERSITY • 30 tests from 20 different experiments • Mesocosm and exclosure experiments • 𝒳2 P = 0.01 • Intuitive result, yet rarely tested experimentally • In cases where diversity of a higher trophic level was positively correlated with a lower trophic level, the higher trophic level had other functions • Seed dispersal, habitat creation Negative effect Positive effect NS Lower trophic level measured 4 2 2 Higher trophic level measured 0 13 2
  • 21. CONCLUSIONS • Herbivore diversity does not beget woody diversity • Herbivory ≠ Herbivore diversity; lots of individual interactions make up broader diversity patterns and relationships • Need more studies that manipulate multiple levels of diversity • Incorporating trophic interactions gives us a better understanding of biodiversity patterns
  • 22. KLEE: • TrumanYoung • KariVeblen • Corinna Riginos • Duncan Kimuyu Smithsonian: • David Kenfack • Paul Musili Field assistance: • Mathew Namoni • Jackson Ekadeli Thanks!