Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna Heather Campbell, Mark Fellowes & James Cook School of Biological Sciences, University of Reading, Reading, RG6 6AS

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) Ants and Acacias Ant–plant associations are widespread (in over 100 genera of tropical angiosperms and at least 20 plant families and in five of 12 ant subfamilies) Research focus on 3 plant genera: Acacia; Cecropia; and Macaranga Ant benefits: Shelter Food - extrafloral nectar, food bodies, third party insects i.e. aphids, scale insects Acacia benefits: Reduced encroachment – competition & fire break Pollination Protection against herbivores

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) Acacia-dominated savanna, Kuzikus Wildlife Reserve, Namibian Central Kalahari Study site Kuzikus is approximately 180km south east of Windhoek, situated in the Namibian Central Kalahari. The study habitat is Acacia-dominated savannah; however the site also incorporates dunes, saltpans, bushveldt and grassland.

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) Insects e.g. Gonometa postica caterpillars Acacia erioloba Species Pollinators Thorn-dwelling ants Herbivores Mammals TetraponeraCrematogaster Cataulacus Tapinoma Sap sucking insects

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) Regarding herbivory on A. erioloba we tested: 1. Do patrolling worker ants act as a deterrent to insect herbivores? 2. How important are G. postica larvae in the defoliation of trees? 3. Which ant species are responsible for antiherbivore activity? Questions

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) 32 trees sampled (16 trees per trial) – two adjacent branches selected; ants excluded from experimental branch and control left with natural ant fauna After 11 days branches removed with secateurs and sealed in plastic bags – this captured any herbivores present on the branch Estimates of leaf damage on each branch made by recording the number of leaflets with herbivore damage out of the total number of leaflets (for 15 leaves per branch) Herbivores collected from branches were counted and identified Experiment repeated twice – before and after hatching of Gonometa silk moth larvae Methods

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) Results 1. Do patrolling worker ants act as a deterrent to insect herbivores? The mean (+SE) percentage of leaflets damaged per branch, according to treatment, for the first and second trials of the ant exclusion experiment. First trial; damage was 13% on ant excluded branches, nearly double (7%) the damage on branches with patrolling ants (t 16 = -3.52, p<0.005). Second trial ; branches without ants had damage to 33% of their leaves as opposed to 11% damage in branches with ants (t 16 =-6.73, p<0.001). N = 16 trees for both trials. Error bars show standard errors. Branches without patrolling ants showed herbivore damage to 22% of leaflets, whereas branches with ants had only 9% leaflet damage (t 32 = -6.83, p < 0.001).

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) Results 1. Do patrolling worker ants act as a deterrent to insect herbivores? Herbivorous insects were found in greater numbers on branches without ants than on those with ants (t 32 = , p < 0.001), although the relationship was more pronounced in the second experiment than in the first (first experiment; t 16 = -2.42, p < 0.05; second experiment t 16 = -3.04, p < 0.01). The mean (+SE) number of insects per branch, according to treatment, for the first and second trials collected at the end of the ant exclusion experiment. N = 16 trees for both trials. Error bars show standard errors.

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) Results 2. How important are G. postica larvae in the defoliation of A. erioloba trees? The number of hemipterans and coleopterans are significantly lower on ant patrolled branches than branches without ants (Hemiptera: t 32 = -3.33, p < 0.005, Coleoptera: t 32 = 2.31, p<0.05). The number of lepidopteran larvae are not affected by the presence or absence of patrolling ants (t 32 = , p = ). There was no increase in lepidopteran numbers between trials on control or experimental branches (control: t 16 = 1.57, ns, experimental: t 16 =1.31, ns). So differences in levels of herbivory cannot be attributed to an increase in presence of Gonometa larvae.

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) Results 3. Which ant species on A. erioloba are responsible for antiherbivore activity? The average percentage of leaflets damaged per branch, for branches with patrolling ants, according to ant species present. The category of multiple ants represents any combination of ant species where more than one species was present. Damage was recorded as number of herbivore damaged leaflets out of total number of leaflets per pinna. Pinna data was pooled for all leaves measured for damage on each branch, giving an average of leaflet damage per branch. N = 16 trees. Analysis showed no significant association between herbivore damage to leaflets and different ant species present on branches (ANOVA, n=32, d.f.=4, ns). Behavioural experiments suggest only Crematogaster behave aggressively, which supports data

Interactions between domatia-dwelling ants & Acacia erioloba in Namibian savanna (Campbell et al 2012) Conclusion Ants DO defend A. erioloba but are mainly effective against hemipteran herbivores Further research needed to identify which species are responsible for defence Tetraponera Cataulacus CrematogasterTapinoma

 BBSRC for funding my PhD  Ministry of Environment and Tourism, Namibia for permission to carry out this research (permit # 1614/2011)  My supervisors Dr Mark Fellowes and Professor James Cook  My field assistant Ian Townsend  Staff at Kuzikus for logistical support  Dr Graham Stone, University of Edinburgh for advice and loan of equipment  Brian Taylor for advice on ant identification Acknowledgements