TheThe influenceinfluence ofof herbivoryherbivory onon grasslandgrassland communitycommunity compositioncomposition andand diversitydiversity inin thethe KrugerKruger NationalNational Park:Park: disentanglingdisentangling thethe rolesroles ofof herbherbivoreivore sizesize andand habitathabitat fertilityfertility

Richard W.S. Fynn, Deron E. Burkepile, Alan K. Knapp, Melinda D. Smith, & Navashni Govender

Savanna Convergence Experiment

University of Kwazulu-Natal, Colorado State University, Yale University, Kansas State University, University of New Mexico, Kruger National Park AreAre allall herbivoresherbivores thethe same?same?

DoesDoes herbivoreherbivore richnessrichness matter?matter? TheThe AquariusAquarius

HERBIVOREHERBIVORE SPECIESSPECIES RICHNESSRICHNESS ISIS CRUCIALCRUCIAL TOTO REEFREEF ECOSYSTEMECOSYSTEM FUNCTION!FUNCTION! GrazingGrazing EcosystemsEcosystems DoDo thethe bigbig guysguys runrun thethe show?show? IncreasingIncreasing HumanHuman EncroachmentEncroachment SimulatingSimulating ConsumerConsumer LossLoss

1. Open Areas – No barrier Allows - Elephant, , Rhino, Buffalo, , Zebra, , , , ,

2. Tall Partial Exclosure - Wire fence beginning at 1.8 m Allows – Rhino, Buffalo, Wildebeest, Zebra, Kudu, Impala, Warthog, Steenbok, Duiker

Excludes – Elephant, Giraffe

3. Short Partial Exclosure - Wire fence beginning at 0.9m Allows - Impala, Warthog, Steenbok, Duiker

Excludes - Elephant, Giraffe, Rhino, Buffalo, Wildebeest, Zebra, Kudu

4. Full Exclosure – Diamond Mesh Fence from 0 - 1.2 m Excludes - All Herbivores >0.5 kg EffectsEffects ofof differentdifferent sizesize--classesclasses

AllAll herbivoresherbivores == FullFull ExclosureExclosure vs.vs. OpenOpen AreaArea

LargeLarge herbivoresherbivores == TallTall PartialPartial ExclosureExclosure vs.vs. OpenOpen AreaArea

MediumMedium herbivoresherbivores == TallTall PartialPartial ExclosureExclosure vs.vs. ShortShort PartialPartial ExclosureExclosure

SmallSmall herbivoresherbivores == ShortShort PartialPartial ExclosureExclosure vs.vs. FullFull ExclosureExclosure TallTall PartialPartial ExclosureExclosure

1.8 m ShortShort PartialPartial ExclosureExclosure

0.9 m

ExperimentalExperimental DesignDesign •• ExperimentalExperimental BurnBurn PlotsPlots ((EBP’sEBP’s)) SataraSatara RegionRegion •• Annual,Annual, Triennial,Triennial, UnburnedUnburned PlotsPlots –– nn == 22 •• NestedNested experimentalexperimental designdesign •• nn == 55 forfor eacheach ExclosureExclosure TreatmentTreatment perper burnburn plotplot ResponseResponse MetricsMetrics •• PlantPlant speciesspecies cover,cover, richness,richness, andand diversitydiversity •• PrimaryPrimary productionproduction •• LightLight availabilityavailability •• TreeTree seedlingseedling survivorshipsurvivorship andand growthgrowth TreatmentTreatment EffectivenessEffectiveness •• SpoorSpoor monitoringmonitoring •• DungDung countscounts •• MotionMotion--activatedactivated camerascameras EffectivenessEffectiveness ofof PartialPartial ExclosuresExclosures

Impala Steenbok Wildebeest 15 Zebra Giraffe Elephant

10

5 Tracks/Plot/Survery

closures x Open Areas

Tall Partial E Short Partial Exclosures ImpactImpact andand ContributionContribution

•• ExaminesExamines aa realisticrealistic patternpattern ofof consumerconsumer lossloss fromfrom savannasavanna ecosystemsecosystems

•• BuildsBuilds onon similarsimilar studiesstudies fromfrom HluhluweHluhluwe (Bond(Bond andand OlffOlff),), KenyaKenya (Young(Young etet al.),al.), andand KNPKNP (River/Savanna(River/Savanna Boundaries)Boundaries) examiningexamining effectseffects ofof differentdifferent herbivoresherbivores atat communitycommunity andand ecosystemecosystem levellevel

•• MarineMarine vs.vs. TerrestrialTerrestrial GrazingGrazing Ecosystems:Ecosystems: AreAre ParrotfishParrotfish JustJust WetWet Wildebeest?Wildebeest? ContextContext--dependantdependant effectseffects ofof herbivory:herbivory: thethe importanceimportance ofof habitathabitat productivityproductivity andand herbivoreherbivore sizesize Redrawn from Proulx & Mazumder 1998 (Ecology 79:2581-2592)

30

Low grazing intensity High grazing intensity

20

10 Plant species richness

0 Nutrient-poor Nutrient-rich systems systems Redrawn from Bakker et al. 2006 (Ecology Letters 9:780-788)

Effect of large and small herbivores 80

60 P = 0.003

40

20

0

-20

-40

-60 0 100 200 300 400 500 600

Effect of small herbivores 60 P = 0.45 40

Change in species richness (%) 20

0

-20

-40

-60 0 100 200 300 400 500 600 Primary production (g m-2) Redrawn from Osem et al. 2002 (J. Ecology 90:936-946)

Productivity 500

) Poorly-drained -2 400 Well-drained

300

200

100 Dry Biomass (g m 0

Poorly-drained site

14

12

10 Richness

8

6 Ungrazed Grazed Well-drained site 11

10

9

8

7 Richness

6

5 1996 1997 1998 1999 Year Adapted from Grime (1973)

Stress Optimal Competitive tolerance coexistence exclusion

species richness Light availability Species richness

Environmental stress Habitat productivity Light availability Light availability

Competition intensity Competition intensity

Grazing intensity Productive habitats

Unproductive habitats Stress on plant

Grazing intensity Adapted from Grime (1973)

Stress Optimal Competitive tolerance coexistence exclusion

species richness Light availability Species richness

Environmental stress Habitat productivity Light availability Light availability

Competition intensity Competition intensity

Grazing intensity

AcknowledgementsAcknowledgements

Data collection and Logistics: Beth Amendola Greg Buis Allison Burkepile Catherine Burns Annikki Chamberlain Scott Collins Dave Hoover Kevin Kirkman Dave Thompson

Fencing Crew: Thembi, Wisani, Justice, Conright, Difference

Funding: National Science Foundation, U.S.A.