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Squirrels:
Predators or Dispersers of Marula Nuts?




             Emma Alterman and Peter Macfarlane
               Organization for Tropical Studies
                         May 2, 2011
Introduction         Methods          Results       Discussion


                          Marula
               (Sclerocarya birrea subsp. caffra)
Introduction       Methods          Results   Discussion



Dispersal
                       Bottlenecks

                                                        Adult
                                                       Mortality

                        Fire Trap/
                         Browse

       Establishment




                                 Time
Introduction       Methods          Results   Discussion



Dispersal
                       Bottlenecks

                                                        Adult
                                                       Mortality

                        Fire Trap/
                         Browse

       Establishment




                                 Time
Introduction   Methods             Results        Discussion


                    Tree Squirrels
               (Paraxerus cepapi subsp. cepapi)
Introduction    Methods          Results       Discussion



              Janzen-Connell Hypothesis




I=SeedsShadow
P=Probability of seedling survival because of lower predation
Introduction   Methods   Results   Discussion



                 Our Goals
Introduction     Methods             Results   Discussion



                       The Set Up




        “Near”             “Intermediate”              “Far”
Introduction   Methods     Results   Discussion


                  In the open



                  Under trees




                  In the bush
Introduction   Methods     Results   Discussion




                         •Eaten
                         •Scatter-hoarded
                         •Moved
                         •Missing
Introduction     Methods   Results   Discussion


               Scatter-hoarding
Introduction     Methods   Results   Discussion

UV Torch
                    Scatter-hoarding
Introduction                         Methods            Results   Discussion

                         12

 Number of Marula Nuts
                         10




                          8


                                                                   Untouched
                          6
                                                                   Eaten

                          4
                                                                   Scatter-Hoarded

                                                                   Missing
                          2




                          0

                              Near             Intermediate       Far


 Predation seen directly under the marula trees
 Mix of eating, scatter-hoarding and unknowns
Introduction                                           Methods           Results          Discussion


                                                 10




          Number of Marula Nuts left untouched
                                                  9

                                                  8

                                                  7

                                                  6

                                                  5

                                                  4

                                                  3

                                                  2

                                                  1

                                                  0


                                                      Under Marula Trees      Under Other Trees




Similar predation rates under marula and non-marula trees
Introduction                                      Methods       Results           Discussion


                                             10




      Number of Marula Nuts left untouched
                                              9


                                              8


                                              7


                                              6


                                              5


                                              4


                                              3


                                              2


                                              1


                                              0


                                                  In the Bush        On the Golf Course




 Lower predation rates in the bush than on the golf course
Introduction                               Methods               Results             Discussion

                        40


                        35


                        30


                        25


                        20


                        15


                        10
      Number of Marula Nuts




                              5


                              0
                                  -4   0      4      8     12      16      20   24

                                           Distance of Scatter-Hoarding (m)



 Most nuts were scatter-hoarded close to the tree
Introduction         Methods                 Results   Discussion



         Janzen-Connell Hypothesis




                 Critical Point?

        “Near”                     “Intermediate”              “Far”
Introduction         Methods                Results               Discussion


                    Under Trees                          In The Open
               Predation under marula and                 Fear predation
                 non-marula trees same                Feel safest in the trees
                    -Trees as triggers
Introduction         Methods                  Results               Discussion


                    Under Trees                            In The Open
               Predation under marula and                   Fear predation
                 non-marula trees same                  Feel safest in the trees
                    -Trees as triggers



                                            But…

                                      In The Bush
                               To much cover might make it
                                difficult for squirrels to find
                                              nuts
Introduction   Methods   Results   Discussion
Elephants!
Introduction             Methods      Results   Discussion



        Can travel 6km per day

     Food in gut for up to 46 hours

Excreted food deposited far from source

     Piles good foraging locations

   Can be filled with up to 700 nuts
Introduction   Methods   Results   Discussion
Introduction   Methods    Results            Discussion




                              Nut found by squirrel

                         Scatter-hoarded near dung pile

                         Nut buried far away from parent
                                      trees!
Introduction            Methods      Results   Discussion


     Why is this an effective
      dispersal method?
Seed mortality in elephant gut very low
    • Seed can travel far safely


Buried seeds protected from threats
     • Risk of being burned
     • Predation
Conclusions   •Janzen-Connell holding true
              for savannas

              •Trees as triggers

              •Marulas possibly dispersed
              by 2 modes of zoochory
Thank you!
Thank you!
             Questions?
References
•   Alterman, E. B. Lind, and A. Martinez. 2011. Where are my nuts? The role of elephants and squirrels in marula (Sclerocarya birrea subsp. caffra) seed
    dispersal. Organization for Tropical Studies report spring 2011. Skukuza, Mpumalanga Province, South Africa.
•   Butchart, D. 2009. Wildlife of South Africa | A Photographic Guide. Struik Nature, Cape Town, SA.
•   Davis, S. 2007. Endozoochory in Subtropical Thicket: comparing effects of species with different digestive systems on seed fate. MSc Thesis, Nelson
    Mandela Metropolitan University, South Africa.
•   Forget, P.M. 1990. Seed dispersal of Vouacapoua Americana (Caesalpiniaceae) by caviomorph rodents in French Guiana. Journal of Tropical Ecology
    6:459-468.
•   Forget, P.M. and S.B. Vander Wall. 2001. Scatter-hoarding rodents and marsupials: convergent evolution on diverging continents. Trends in Ecology &
    Evolution. 16(2):65-67.
•   Gallaher, K.S.L. 2010. Marula (Sclerocarya Birrea Subsp. Caffra) Dispersal by Mammals: Are Squirrels Seed Predators Or Seed Dispersers? BSc. Honors
    Thesis. Botany Department University of Cape Town, Cape Town, SA.
•   Helm, C.V., S.L. Scott, and E.T.F. Witkowski. 2011. Reproductive potential and seed fate of Sclerocarya birrea subsp. caffra (marula) in the low altitude
    savannas of South Africa. South African Journal of Botany. in press.
•   Jacobs, O.S. and R. Biggs. 2002. The status and population structure of the marula in the Kruger National Park. South African Journal of Wildlife
    Research 32(1): 1–12.
•   Janzen, D. 1970. Herbivores and the Number of Tree Species in Tropical Forests. The American Naturalist 104(940):501-528.
•   Laorie, S.R., Van Aarde, R.J., and S.L. Pimm. 2009. Fences and artificial water affect African savannah elephant movement patterns. Biological
    Conservation 142:3086–3098
•   Midgley, J.J. and W.J. Bond. 2001. A synthesis of the demography of African acacias. Journal of Tropical Ecology 17:871–886.
•   Rees, P. A. 1982. Gross assimilation efficiency and food passage time in the African elephant. African Journal of Ecology 20(3):193-198.
•   Shackleton, C.M., J. Botha, P.L. Emanuel and S. Ndlovu. 2002. Inventory of Marula (Sclerocarya birrea subsp. caffra) Stocks and Fruit Yields in
    Communal and Protected Areas of the Bushbuckridge Lowveld, Limpopo Province, South Africa. Department of Environmental Science, Rhodes
    University, Grahamstown 6140, South Africa.
•   Schmidt E., M. Lötter, W. McCleland. 2002. Trees and Shrubs of Mpumalanga and Kruger National Park. Jacana, Johannesburg, SA.
•    Stuart, C. and T. Stuart. 2007 Field Guide to Mammals of Southern Africa. Fourth Edition Struik Piblishers, Cape Town, SA.
•    van Wilgen, B.W., W.S.W. Trollope, H.C. Biggs, A.L.F. Potgieter, and B.H. Brockett. 2003. Fire as a Driver of Ecosystem Variability. Pp 149-170 in J.T.
    du Toit, K.H. Rogers, and H.C. Biggs, editors. The Kruger Experience Ecology and Management of Savanna Heterogeneity. Island Press, Washington,
    DC, USA.

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Marula Dispersal

  • 1. Squirrels: Predators or Dispersers of Marula Nuts? Emma Alterman and Peter Macfarlane Organization for Tropical Studies May 2, 2011
  • 2. Introduction Methods Results Discussion Marula (Sclerocarya birrea subsp. caffra)
  • 3. Introduction Methods Results Discussion Dispersal Bottlenecks Adult Mortality Fire Trap/ Browse Establishment Time
  • 4. Introduction Methods Results Discussion Dispersal Bottlenecks Adult Mortality Fire Trap/ Browse Establishment Time
  • 5. Introduction Methods Results Discussion Tree Squirrels (Paraxerus cepapi subsp. cepapi)
  • 6. Introduction Methods Results Discussion Janzen-Connell Hypothesis I=SeedsShadow P=Probability of seedling survival because of lower predation
  • 7. Introduction Methods Results Discussion Our Goals
  • 8. Introduction Methods Results Discussion The Set Up “Near” “Intermediate” “Far”
  • 9. Introduction Methods Results Discussion In the open Under trees In the bush
  • 10. Introduction Methods Results Discussion •Eaten •Scatter-hoarded •Moved •Missing
  • 11. Introduction Methods Results Discussion Scatter-hoarding
  • 12. Introduction Methods Results Discussion UV Torch Scatter-hoarding
  • 13. Introduction Methods Results Discussion 12 Number of Marula Nuts 10 8 Untouched 6 Eaten 4 Scatter-Hoarded Missing 2 0 Near Intermediate Far Predation seen directly under the marula trees Mix of eating, scatter-hoarding and unknowns
  • 14. Introduction Methods Results Discussion 10 Number of Marula Nuts left untouched 9 8 7 6 5 4 3 2 1 0 Under Marula Trees Under Other Trees Similar predation rates under marula and non-marula trees
  • 15. Introduction Methods Results Discussion 10 Number of Marula Nuts left untouched 9 8 7 6 5 4 3 2 1 0 In the Bush On the Golf Course Lower predation rates in the bush than on the golf course
  • 16. Introduction Methods Results Discussion 40 35 30 25 20 15 10 Number of Marula Nuts 5 0 -4 0 4 8 12 16 20 24 Distance of Scatter-Hoarding (m) Most nuts were scatter-hoarded close to the tree
  • 17. Introduction Methods Results Discussion Janzen-Connell Hypothesis Critical Point? “Near” “Intermediate” “Far”
  • 18. Introduction Methods Results Discussion Under Trees In The Open Predation under marula and Fear predation non-marula trees same Feel safest in the trees -Trees as triggers
  • 19. Introduction Methods Results Discussion Under Trees In The Open Predation under marula and Fear predation non-marula trees same Feel safest in the trees -Trees as triggers But… In The Bush To much cover might make it difficult for squirrels to find nuts
  • 20. Introduction Methods Results Discussion
  • 22. Introduction Methods Results Discussion Can travel 6km per day Food in gut for up to 46 hours Excreted food deposited far from source Piles good foraging locations Can be filled with up to 700 nuts
  • 23. Introduction Methods Results Discussion
  • 24. Introduction Methods Results Discussion Nut found by squirrel Scatter-hoarded near dung pile Nut buried far away from parent trees!
  • 25. Introduction Methods Results Discussion Why is this an effective dispersal method? Seed mortality in elephant gut very low • Seed can travel far safely Buried seeds protected from threats • Risk of being burned • Predation
  • 26. Conclusions •Janzen-Connell holding true for savannas •Trees as triggers •Marulas possibly dispersed by 2 modes of zoochory
  • 28. Thank you! Questions?
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