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Effects of Fire and Herbivory 
on Tree Size Transitions in 
Acacia drepanolobium 
Eric M LaMalfa 
Preliminary data 
11/6/2014
Tree cover in savannas 
Top-down controls on tree cover 
• Disturbance: Fire AND/OR Herbivory 
Are tree size transitions controlled by herbivory x fire?
Fire effects: 
•Topkill 
•Influences plant-plant intxns 
- reduced competition for resources
Grazing effects: 
•Reduces herbaceous biomass 
•Influences plant-plant intxns 
- Reduced competition for 
1) unpalatable herbaceous plants 
2) woody plants
Browsing effects 
•Reduced woody biomass 
•Reduced seedling establishment 
•Influences plant-plant intxns 
- Reduced competition for herbaceous 
plants
Direct fire x herbivory intxns 
• Herbivore preference for burned areas 
 Increased crude protein content 
 Decreased lignin
Direct fire x herbivory intxns 
• Grazing reduces fine fuel load/fire continuity 
Increased tree survival 
• Browsing reduces heavy fuel and fire temps 
Increased tree survival
Indirect fire x herbivory intxns 
Fire kills tree-obligate ants 
Decreased ant-defense 
Increased browse pressure 
Reduced tree fitness
Laikipia plateau, Kenya 
• 1800m 
• Semi-arid 550-600 mm 
Cattle 
Wildife 
Mega-herbivores
Acacia 
drepanolobium 
• 95% of tree cover 
• Nitrogen fixing bacteria 
nodules 
• Highly quality (browse and 
herbaceous) 
• Coppices (re-sprouts after fire 
and browse damage)
Trees Provide 
Nest & Food 
Crematogaster sjostedt 
C. mimosae 
C. nigriceps 
Tetraponera penzigi 
Extra Floral 
Nectary 
Domatia 
Ants Provide 
Plant Defense 
• High defense against herbivores 
and insects 
• High nectar use 
• High resistance to fire
• Ants track tree resources during sapling colonization 
• Trees > 1m (grass height) are rarely observed without ants
Research Questions 
1) How do fire and herbivory affect small tree (<1m) 
size transitions? 
2) How do fire and herbivory affect ant colonization 
of small trees?
Experimental Design – Herbivory 
• Kenya Long Term Exclosure Experiment (KLEE) 
• Fences exclude herbivores (since 1995) 
• Randomized complete block design 
MW & MWC 
W & WC 
0 & C 
C= cattle 
W= wildlife 
M= mega-herbivores 
0= no herbivores 
MWC 
MWC 
MW 
MW 
MWC 
MW 
W 
W 
W 
C 
C 
C 
0 
0 
0 
WC 
WC 
WC
Experimental Design – Fire 
• 18 burns, February 2013 
• 12 treatments, 3 replicates 
• 4304 tagged trees 
• ant sp., tree size, herbivory 
• % top-kill 
Kimuyu D.M., Sensenig R.L., Riginos C., Veblen K.E., and T.P. Young (2014). “Native and domestic browsers and grazers reduce fuels, 
fire temperatures, and acacia ant mortality in an African savanna.” Ecological Applications 24(4): 741–749
Post-Fire Trees <1m 
1 meter 
• 4304 tagged trees 
• ant sp., tree size, herbivory 
• % top-kill
Control (no burn) 
1 3 5 
5 m 
30 m 
Post fire 
N = 36 plots 
27 to 190 trees/ plot 
1659 trees <1m height
height 
Main stem length 
Basal stem length 
Branch stem length 
Total stem length 
Green stem length 
Main stem diameter 
Domatia diameter 
Green stem diameter 
Domatia count 
Basal stem count 
Green stem count 
Nectary count (5cm) 
Ant species 
Data 
5cm
PRELIMINARY RESULTS 
How do fire and herbivory affect small 
tree (<1m) size transitions? 
1m 
Un-burned Burned 
N = 666 N = 993
5cm 
Tree size (<1m) 
39 25 233 254 
t = -1.6733, 
df = 1292.361, 
p-value = 0.09451 
t = 16.1482, 
df = 1013.013, 
p-value < 2.2e-16 
Tree Height Total Stem Length
Ant mutualism rewards (domatia) 
t = 3.2196, 
df = 1000.367, 
p-value = 0.001325 
9.95 7.63 
t = -1.7532, 
df = 1255.261, 
p-value = 0.07981 
9.9 10.5
Ant mutualism rewards (nectar) 
0.61 1.24 
t = -7.5473, 
df = 1654.985, 
p-value = 7.295e-14 
12 31 
t = -10.0173, 
df = 1598.088, 
p-value < 2.2e-16 
0.74 1.33 
t = -13.8396, 
df = 1618.421, 
p-value < 2.2e-16 
1.73 3.33 
t = -9.2696, 
df = 1645.348, 
p-value < 2.2e-16
Preliminary results 
How do fire and herbivory affect ant 
colonization of small trees? 
Extra Floral Nectary
Green Stem Length 
Un-burned Burned
Green Stem Diameter 
Un-burned Burned
Nectary Count 
Un-burned Burned
Conclusions 
1. Mean tree size in the <1m size class is 
no different 1.5 years following fire. 
2. Ant mutualism traits associated with 
nectar are higher in burned trees. 
3. After fire, small trees occupied by C. 
nigriceps (BBR) and C. mimosae (RRB) 
& have higher nectar rewards.
Do large top-killed trees produce 
better nectar rewards? 
1 meter 
Next Steps
Next Steps 
Does fire temperature affect tree response? 
Low heat > High heat 
Does herbivory affect tree response? 
Low browse > High browse
Next Steps 
Does neighbor tree colony affect post fire 
ant colonization? 
Neighbor Data 
ant 
damage 
neighbor_tree_1 
neighbor_tree_2 
neighbor_tree_3 
neighbor_tree_4 
neighbor_tree_5 
neighbor_tree_6 
neighbor_ant 
2m
Thanks 
S.J. and Jessie E. 
Quinney 
Foundation 
Kari Veblen PhD - Advisor 
Mellisa Harvey LaMalfa – Mutualism 
Veblen Lab – Rebecca, Kyle, Maike
Correlation matrix 
ms_length 
tot_length 
green_length 
main_diam 
domatia_diam 
green_diam 
domatia_count 
basal_count 
green_count 
nectary_count 
damage 
height 0.78 0.54 0.13 0.72 0.50 0.07 0.59 -0.14 0.10 0.14 -0.03 
ms_length 0.56 0.14 0.72 0.55 0.11 0.59 -0.15 0.11 0.16 -0.07 
tot_length 0.48 0.62 0.54 0.37 0.74 0.36 0.50 0.34 0.08 
green_length 0.22 0.30 0.50 0.35 0.34 0.80 0.47 0.13 
main_diam 0.56 0.21 0.67 -0.07 0.25 0.24 0.08 
domatia_diam 0.36 0.57 0.14 0.28 0.32 0.15 
green_diam 0.29 0.25 0.52 0.50 0.20 
domatia_count 0.08 0.33 0.33 0.03 
basal_count 0.37 0.18 0.19 
green_count 0.44 0.15 
nectary_count 0.08

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Effects of Fire and Herbivory on Tree Size and Ant Colonization in Acacia Savanna

  • 1. Effects of Fire and Herbivory on Tree Size Transitions in Acacia drepanolobium Eric M LaMalfa Preliminary data 11/6/2014
  • 2. Tree cover in savannas Top-down controls on tree cover • Disturbance: Fire AND/OR Herbivory Are tree size transitions controlled by herbivory x fire?
  • 3. Fire effects: •Topkill •Influences plant-plant intxns - reduced competition for resources
  • 4. Grazing effects: •Reduces herbaceous biomass •Influences plant-plant intxns - Reduced competition for 1) unpalatable herbaceous plants 2) woody plants
  • 5. Browsing effects •Reduced woody biomass •Reduced seedling establishment •Influences plant-plant intxns - Reduced competition for herbaceous plants
  • 6. Direct fire x herbivory intxns • Herbivore preference for burned areas  Increased crude protein content  Decreased lignin
  • 7. Direct fire x herbivory intxns • Grazing reduces fine fuel load/fire continuity Increased tree survival • Browsing reduces heavy fuel and fire temps Increased tree survival
  • 8. Indirect fire x herbivory intxns Fire kills tree-obligate ants Decreased ant-defense Increased browse pressure Reduced tree fitness
  • 9. Laikipia plateau, Kenya • 1800m • Semi-arid 550-600 mm Cattle Wildife Mega-herbivores
  • 10. Acacia drepanolobium • 95% of tree cover • Nitrogen fixing bacteria nodules • Highly quality (browse and herbaceous) • Coppices (re-sprouts after fire and browse damage)
  • 11. Trees Provide Nest & Food Crematogaster sjostedt C. mimosae C. nigriceps Tetraponera penzigi Extra Floral Nectary Domatia Ants Provide Plant Defense • High defense against herbivores and insects • High nectar use • High resistance to fire
  • 12. • Ants track tree resources during sapling colonization • Trees > 1m (grass height) are rarely observed without ants
  • 13. Research Questions 1) How do fire and herbivory affect small tree (<1m) size transitions? 2) How do fire and herbivory affect ant colonization of small trees?
  • 14. Experimental Design – Herbivory • Kenya Long Term Exclosure Experiment (KLEE) • Fences exclude herbivores (since 1995) • Randomized complete block design MW & MWC W & WC 0 & C C= cattle W= wildlife M= mega-herbivores 0= no herbivores MWC MWC MW MW MWC MW W W W C C C 0 0 0 WC WC WC
  • 15. Experimental Design – Fire • 18 burns, February 2013 • 12 treatments, 3 replicates • 4304 tagged trees • ant sp., tree size, herbivory • % top-kill Kimuyu D.M., Sensenig R.L., Riginos C., Veblen K.E., and T.P. Young (2014). “Native and domestic browsers and grazers reduce fuels, fire temperatures, and acacia ant mortality in an African savanna.” Ecological Applications 24(4): 741–749
  • 16. Post-Fire Trees <1m 1 meter • 4304 tagged trees • ant sp., tree size, herbivory • % top-kill
  • 17. Control (no burn) 1 3 5 5 m 30 m Post fire N = 36 plots 27 to 190 trees/ plot 1659 trees <1m height
  • 18. height Main stem length Basal stem length Branch stem length Total stem length Green stem length Main stem diameter Domatia diameter Green stem diameter Domatia count Basal stem count Green stem count Nectary count (5cm) Ant species Data 5cm
  • 19. PRELIMINARY RESULTS How do fire and herbivory affect small tree (<1m) size transitions? 1m Un-burned Burned N = 666 N = 993
  • 20. 5cm Tree size (<1m) 39 25 233 254 t = -1.6733, df = 1292.361, p-value = 0.09451 t = 16.1482, df = 1013.013, p-value < 2.2e-16 Tree Height Total Stem Length
  • 21. Ant mutualism rewards (domatia) t = 3.2196, df = 1000.367, p-value = 0.001325 9.95 7.63 t = -1.7532, df = 1255.261, p-value = 0.07981 9.9 10.5
  • 22. Ant mutualism rewards (nectar) 0.61 1.24 t = -7.5473, df = 1654.985, p-value = 7.295e-14 12 31 t = -10.0173, df = 1598.088, p-value < 2.2e-16 0.74 1.33 t = -13.8396, df = 1618.421, p-value < 2.2e-16 1.73 3.33 t = -9.2696, df = 1645.348, p-value < 2.2e-16
  • 23. Preliminary results How do fire and herbivory affect ant colonization of small trees? Extra Floral Nectary
  • 24. Green Stem Length Un-burned Burned
  • 25. Green Stem Diameter Un-burned Burned
  • 27. Conclusions 1. Mean tree size in the <1m size class is no different 1.5 years following fire. 2. Ant mutualism traits associated with nectar are higher in burned trees. 3. After fire, small trees occupied by C. nigriceps (BBR) and C. mimosae (RRB) & have higher nectar rewards.
  • 28. Do large top-killed trees produce better nectar rewards? 1 meter Next Steps
  • 29. Next Steps Does fire temperature affect tree response? Low heat > High heat Does herbivory affect tree response? Low browse > High browse
  • 30. Next Steps Does neighbor tree colony affect post fire ant colonization? Neighbor Data ant damage neighbor_tree_1 neighbor_tree_2 neighbor_tree_3 neighbor_tree_4 neighbor_tree_5 neighbor_tree_6 neighbor_ant 2m
  • 31. Thanks S.J. and Jessie E. Quinney Foundation Kari Veblen PhD - Advisor Mellisa Harvey LaMalfa – Mutualism Veblen Lab – Rebecca, Kyle, Maike
  • 32. Correlation matrix ms_length tot_length green_length main_diam domatia_diam green_diam domatia_count basal_count green_count nectary_count damage height 0.78 0.54 0.13 0.72 0.50 0.07 0.59 -0.14 0.10 0.14 -0.03 ms_length 0.56 0.14 0.72 0.55 0.11 0.59 -0.15 0.11 0.16 -0.07 tot_length 0.48 0.62 0.54 0.37 0.74 0.36 0.50 0.34 0.08 green_length 0.22 0.30 0.50 0.35 0.34 0.80 0.47 0.13 main_diam 0.56 0.21 0.67 -0.07 0.25 0.24 0.08 domatia_diam 0.36 0.57 0.14 0.28 0.32 0.15 green_diam 0.29 0.25 0.52 0.50 0.20 domatia_count 0.08 0.33 0.33 0.03 basal_count 0.37 0.18 0.19 green_count 0.44 0.15 nectary_count 0.08

Editor's Notes

  1. Demographic bottleneck – important transition browsers eat trees, persistent short stout trees. Grazers facilitate large tree growth and high density tree canopy. After low temperature fire tall trees survive and thrive Fires are hot and top-kill more trees all at once. Important because these are the primary means with which humans affect these rangelands, Management objectives include tree cover +/-
  2. Ammount of kill vs biomass removal depends on temperature
  3. Selectivity for palatibility
  4. Selectivity for palatability
  5. The level of utilization is effected by fire.
  6. The effects of fire depends on herbivory regime
  7. Elephants avoid trees with these ants. Very effective plant defence.
  8. East Africa……..Equator………rain shadow of Mt. Kenya. Three types of herbivores
  9. Elephants top-kill trees
  10. Point to the screen and say the names of the ant species of interist. Define domatia – swollen thorns Define - Extra floral nectary
  11. Dominant to submissive hierarchy –
  12. Small trees less than one meter (demographic bottleneck) important life stage transition for savannas.
  13. O, M, W, C
  14. Pre- fire data is important because not all coppicing trees are equal.
  15. Quick Post fire 1.5 years
  16. Quick How does fire effect size and mutualism rewards.
  17. Individual trees as sample units Focus on burned vs unburned trees (pooled across herbivore treatments)
  18. Unburned trees are taller No difference in total stem length
  19. There are 20% more domatia per plant, however, the difference is not biologically significant (8-10 domatia) No difference in the size of domatia
  20. Green Stem Diameter 2x in burned trees Green Stem count 2x in burned trees Nectary count 2X in burned trees Green Stem length 2X in burned trees
  21. Looking at the pooled trees again how different are the mutualism rewards on trees occupied by the different ant species (burned vs unburned).
  22. Green stem length is different between burned and unburned -except for unoccupied trees more like unburned Not different among different ant-trees
  23. Green stem diameter is different between burned and unburned -except for unoccupied trees more like unburned -TP large variance possibly due to submissive position and dispersal mechanism Not different among different ant-trees
  24. In burned trees more nectaries per green tip after fire in RRB and BBR Could mean that the trees have better resources that the ants are tracking or the ants are effectively protecting nectaries from being browsed or the presence of ants are modifying the traits
  25. INCREASE MATRIX, GET RID OF THE TABLE ON FAR LEFT BECAUSE IT’S REDUNDANT.