Abstract
Sexual differences in foraging behaviour of African elephant are comparatively well studied. However, the behaviour is known to vary between savannas due to differences in plant and herbivory communities which determine the level of forage selection and competitions. The sexual variation in foraging behaviour is attributed by sexual dimorphism, where male elephants are larger than females. The different body sizes result in divergent foraging strategies due to the differential scaling coefficients of metabolism and gut size, where the larger bodied individuals can tolerate greater quantities of low-quality forage. We studied the foraging behaviour of male and female elephants in Serengeti National Park, a nutrient-rich savanna in Tanzania, based on the forage selection and the scramble competition hypotheses. Data were collected for 2 months in 2014 by driving in road transects and make visual observations of browsing elephants within 50 m from the road. We recorded the number of bites taken per tree, parts of the tree browsed, browsing height, and tree species. Females targeted a wider range of wood species, browsing at lower heights compared to males. The sexes had a comparable length of browsing bouts, spending an average of 4 min per tree. The results support the scramble competition hypothesis through the feeding height stratification and the forage selection hypothesis through the higher diversity of tree species utilised by females. We suggest that sexual differences in forage requirements interact with forage quality and competing herbivore communities in shaping the foraging behaviour of elephants.
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References
Anderson TM, Dempewolf J, Metzger KL, Reed DN, Serneels S (2008) Generation and maintenance of heterogeneity in the Serengeti ecosystem. In: Sinclair ARE, Packer C, Mduma SAR, Fryxell JM (eds) Serengeti III: human impacts on ecosystem dynamics. University of Chicago Press, Chicago, Illinois, pp 135–182
Archie EA, Chiyo PI (2012) Elephant behaviour and conservation: social relationships, the effects of poaching, and genetic tools for management. Mol Ecol 21:765–778. https://doi.org/10.1111/j.1365-294x.2011.05237.x
Arsenault R, Owen-Smith N (2002) Facilitation versus competition in grazing herbivore assemblages. Oikos 97:313–318. https://doi.org/10.1034/j.1600-0706.2002.970301.x
Augustine DJ, Mcnaughton SJ (2004) Regulation of shrub dynamics by native browsing ungulates on East African rangeland. J Appl Ecol 41:45–58. https://doi.org/10.1111/j.1365-2664.2004.00864.x
Barnes RFW (1980) The decline of the baobab tree in Ruaha National Park, Tanzania. Afr J Ecol 18:243–252. https://doi.org/10.1111/j.1365-2028.1980.tb01053.x
Barnes RFW (1982) Elephant feeding behaviour in Ruaha National Park, Tanzania. Afr J Ecol 20:123–136. https://doi.org/10.1111/j.1365-2028.1982.tb00282.x
Bell RHV (1971) A grazing ecosystem in the Serengeti. Sci Am 225:86–93. https://doi.org/10.1038/scientificamerican0771-86
Bell RHV (1982) The effect of soil nutrient availability on community structure in African ecosystems. In: Huntley BJ, Walker BH (eds) Ecology of tropical savannas. Springer Verlag, Berlin, pp 193–216
Biru Y, Bekele A (2012) Food habits of African elephant (Loxodonta africana) in Babile Elephant Sanctuary, Ethiopia. Trop Ecol 53:43–52
Bryant JP, Kuropat PJ, Cooper SM, Frisby K, Owen-Smith N (1989) Resource availability hypothesis of plant antiherbivore defence tested in a South African savanna ecosystem. Nature 340:227–229. https://doi.org/10.1038/340227a0
Bryant JP, Reichardt PB, Clausen TP (1992) Chemically mediated interactions between woody plants and browsing mammals. J Range Manag 45:18–24. https://doi.org/10.2307/4002520
Bukombe J, Smith S, Kija H, Loishooki A, Sumay G, Mwita M, Mwakalebe G, Kihwele E (2018) Fire regulates the abundance of alien plant species around roads and settlements in the Serengeti National Park. Manag Biol Invasions 9:357–367. https://doi.org/10.3391/mbi.2018.9.3.17
Cameron EZ, du Toit JT (2007) Winning by a neck: tall giraffes avoid competing with shorter browsers. Amer Nat 169:130–135. https://doi.org/10.1086/509940
Chiyo PI, Archie EA, Hollister-Smith JA, Lee PC, Poole JH, Moss CJ, Alberts SC (2011) Association patterns of African elephants in all-male groups: the role of age and genetic relatedness. Anim Behav 81:1093–1099. https://doi.org/10.1016/j.anbehav.2011.02.013
Clutton-Brock TH, Iason GR, Guinness FE (1987) Sexual segregation and density-related changes in habitat use in male and female Red deer (Cerrus elaphus). J Zool 211:275–289. https://doi.org/10.1111/j.1469-7998.1987.tb01534.x
Codron J, Lee-Thorp JA, Sponheimer M, Codron D, Grant RC, de Ruiter DJ (2006) Elephant (Loxodonta africana) diets in Kruger National Park, South Africa: spatial and landscape differences. J Mammal 87:27–34. https://doi.org/10.1644/05-MAMM-A-017R1.1
Coe MJ, Cumming DH, Phillipson J (1976) Biomass and production of large African herbivores in relation to rainfall and primary production. Oecologia 22:341–354. https://doi.org/10.1007/bf00345312
Conradt L, Ruckstuhl KE, Neuhaus P (2005) Definitions, hypotheses, models and measures in the study of animal segregation. In: Ruckstuhl KE, Neuhaus P (eds) Sexual segregation in vertebrates: ecology of the two sexes. Cambridge University Press, Cambridge, United Kingdom, pp 11–32
Cooper SM, Owen-Smith N (1986) Effects of plant spinescence on large mammalian herbivores. Oecologia 68:446–455. https://doi.org/10.1007/BF01036753
Dambe LM, Mogotsi K, Odubeng M, Kgosikoma OE (2015) Nutritive value of some important indigenous livestock browse species in semi-arid mixed Mopane bushveld, Botswana. Livest Res Rural Dev 27(10). http://www.lrrd.org/lrrd27/10/mogo27209.htm
De Knegt HJ, Van Langevelde F, Skidmore AK, Delsink A, Slotow R, Henley S, Bucini G, De Boer WF, Coughenour MB, Grant CC (2011) The spatial scaling of habitat selection by African elephants. J Anim Ecol 80:270–281. https://doi.org/10.1111/j.1365-2656.2010.01764.x
Demment MW (1983) Feeding ecology and the evolution of body size of baboons. Afr J Ecol 21:219–233. https://doi.org/10.1111/j.1365-2028.1983.tb00323.x
Demment MW, Van Soest PJ (1985) A nutritional explanation for body-size patterns of ruminant and nonruminant herbivores. Amer Nat 125:641–672. https://doi.org/10.1086/284369
du Toit JT (1990) Feeding-height stratification among African browsing ruminants. Afr J Ecol 28:55–61. https://doi.org/10.1111/j.1365-2028.1990.tb01136.x
du Toit JT (2005) Sex differences in the foraging ecology of large mammalian herbivores. In: Ruckstuhl KE, Neuhaus P (eds) Sexual segregation in vertebrates: ecology of the two sexes. Cambridge: Cambridge University Press. Cambridge University Press, New York, pp 35–52
Dyer C (2014) New names for the African Acacia species in Vachellia and Senegalia. Southern for A J for Sci 76:4. https://doi.org/10.2989/20702620.2014.980090
East R (1984) Rainfall, soil nutrient status and biomass of large African savanna mammals. Afr J Ecol 22:245–270. https://doi.org/10.1111/j.1365-2028.1984.tb00700.x
Estes RD (2012) The behavior guide to African mammals: including hoofed mammals, carnivores, primates. University of California Press, Berkeley
Evans K, Harris S (2012) Sex differences in habitat use by A frican elephants (Loxodonta africana) in the Okavango Delta, Botswana: is size really the deciding factor? Afr J Ecol 50:277–284. https://doi.org/10.1111/j.1365-2028.2012.01322.x
Freeland WJ, Saladin LR (1989) Choice of mixed diets by herbivores: the idiosyncratic effects of plant secondary compounds. Biochem Sys Ecol 17:493–497. https://doi.org/10.1016/0305-1978(89)90029-x
Fritz H, Duncan P, Gordon IJ, Illius AW (2002) Megaherbivores influence trophic guilds structure in African ungulate communities. Oecologia 131:620–625. https://doi.org/10.1007/s00442-002-0919-3
Ginnett TF, Demment MW (1997) Sex differences in giraffe foraging behavior at two spatial scales. Oecologia 110:291–300. https://doi.org/10.1007/s004420050162
Ginnett TF, Demment MW (1999) Sexual segregation by Masai giraffes at two spatial scales. Afr J Ecol 37:93–106. https://doi.org/10.1046/j.1365-2028.1999.00163.x
Gittleman JL, Thompson SD (1988) Energy allocation in mammalian reproduction. Amer Zool 28:863–875. https://doi.org/10.1093/icb/28.3.863
Guy PR (1976) The feeding behaviour of elephant (Loxodonta africana) in the Sengwa area Rhodesia. S Afr J Wildl Res 6:55–63
Hagos MG, Smit GN (2005) Soil enrichment by Acacia mellifera subsp. detinens on nutrient poor sandy soil in a semi-arid southern African savanna. J Arid Environ 61:47–59. https://doi.org/10.1016/j.jaridenv.2004.08.003
Holdo RM (2003) Woody plant damage by African elephants in relation to leaf nutrients in western Zimbabwe. J Trop Ecol 19:189–196. https://doi.org/10.1017/S0266467403003213
Huntley BJ (1982) Southern African savannas. In: Huntley BJ, Walker BH (eds) Ecology of tropical savannas. Springer-Verlag, Berlin, pp 101–119
Jachmann H, Bell RHV (1985) Utilization by elephants of the Brachystegia woodlands of the Kasungu National Park, Malawi. Afr J Ecol 23:245–258. https://doi.org/10.1111/j.1365-2028.1985.tb00955.xCitations
Jager T (1982) Soils of the Serengeti woodlands, Tanzania. Pudoc Wageningen: Netherland
Jarman PJ (1974) The social organisation of antelope in relation to their ecology. Behaviour 48:215–267. https://doi.org/10.1163/156853974X00345
Kennedy AS, Kennedy V (2014) Animals of the Serengeti: and Ngorongoro Conservation Area. Princeton University Press, USA
Kihwele ES, Veldhuis MP, Loishooki A, Hongoa JR, Hopcraft JGC, Olff H, Wolanski E (2021) Upstream land-use negatively affects river flow dynamics in the Serengeti National Park. Ecohydrol Hydrobiol 21:1–12. https://doi.org/10.1016/j.ecohyd.2020.12.004
Kilungu H, Leemans R, Munishi PK, Amelung B (2017) Climate change threatens major tourist attractions and tourism in Serengeti National Park, Tanzania. In: Leal-Filho W, Belay S, Kalangu J, Menas W, Munishi P, Musiyiwa K (eds) Climate change adaptation in Africa. Springer International Publishing, Cham, Switzerland, pp 375–392
Kinahan A, Pimm SL, Van Aarde RJ (2007) Ambient temperature as a determinant of landscape use in the savanna elephant, Loxodonta africana. J Therm Biol 32:47–58. https://doi.org/10.1016/j.jtherbio.2006.09.002
Kyalangalilwa B, Boatwright JS, Daru BH, Maurin O, van der Bank M (2013) Phylogenetic position and revised classification of Acacia sl (Fabaceae: Mimosoideae) in Africa, including new combinations in Vachellia and Senegalia. Bot J Linn Soc 172:500–523. https://doi.org/10.1111/boj.12047
Loarie SR, van Aarde RJ, Pimm SL (2009) Elephant seasonal vegetation preferences across dry and wet savannas. Biol Conserv 142:3099–3107. https://doi.org/10.1016/j.biocon.2009.08.021
Loveridge AJ, Hunt JE, Murindagomo F, Macdonald DW (2006) Influence of drought on predation of elephant (Loxodonta africana) calves by lions (Panthera leo) in an African wooded savannah. J Zool 270:523–530. https://doi.org/10.1111/j.1469-7998.2006.00181.x
Lu H, Wagner HH, Chen X (2007) A contribution diversity approach to evaluate species diversity. Basic Appl Ecol 8:1–12. https://doi.org/10.1016/j.baae.2006.06.004
Magurran AE (2013) Measuring biological diversity. Blackwell Oxford, UK
Mahenya O, Ndjamba JK, Mathisen KM, Skarpe C (2016) Giraffe browsing in response to plant traits. Acta Oecol 75:54–62. https://doi.org/10.1016/j.actao.2016.07.001
Main MB, Weckerly FW, Bleich VC (1996) Sexual segregation in ungulates: new directions for research. J Mammal 77:449–461. https://doi.org/10.2307/1382821
Makhabu SW (2005) Resource partitioning within a browsing guild in a key habitat, the Chobe Riverfront, Botswana. J Trop Ecol 21:641–649. https://doi.org/10.1017/S0266467405002701
McComb K, Moss C, Sayialel S, Baker L (2000) Unusually extensive networks of vocal recognition in African elephants. Anim Behav 59:1103–1109. https://doi.org/10.1006/anbe.2000.1406
McNaughton SJ, Georgiadis NJ (1986) Ecology of African grazing and browsing mammals. Annu Rev Ecol Syst 17:39–65. https://doi.org/10.1146/annurev.es.17.110186.000351
Mole MA, DÁraujo SR, van Aarde RJ, Mitchell D, Fuller A (2018) Savanna elephants maintain homeothermy under African heat. J Comp Physiol 188:889–897. https://doi.org/10.1007/s00360-018-1170-5
Morrison TA, Estes AB, Mduma SAR, Maliti HT, Frederick H, Kija H, Mwita M, Sinclair ARE, Kohi EM (2018) Informing aerial total counts with demographic models: population growth of Serengeti elephants not explained purely by demography. Conserv Lett 11:e12413. https://doi.org/10.1111/conl.12413
Morrison TA, Holdo RM, Anderson TM (2016) Elephant damage, not fire or rainfall, explains mortality of overstorey trees in Serengeti. J Trop Ecol 104:409–418. https://doi.org/10.1111/1365-2745.12517
Mramba RP, Andreassen HP, Skarpe C (2017a) Browsing and plant traits in nutrient-rich and nutrient-poor savannas in Tanzania. J Trop Ecol 33:327–336. https://doi.org/10.1017/S026646741700030X
Mramba RP, Mahenya O, Siyaya A, Mathisen KM, Andreassen HP, Skarpe C (2017b) Sexual segregation in foraging giraffe. Acta Oecol 79:26–35. https://doi.org/10.1016/j.actao.2016.12.007
Mysterud A (2000) The relationship between ecological segregation and sexual body size dimorphism in large herbivores. Oecologia 124:40–54. https://doi.org/10.1007/s004420050023
O'Kane CAJ, Duffy KJ, Page BR, Macdonald DW (2011) Overlap and seasonal shifts in use of woody plant species amongst a guild of savanna browsers. J Trop Ecol 27:249–258. https://doi.org/10.1017/S0266467410000817
Owen-Smith N, Chafota J (2012) Selective feeding by a megaherbivore, the African elephant (Loxodonta africana). J Mammal 93:698–705. https://doi.org/10.1644/11-mamm-a-350.1
Owen-Smith RN (1992) Megaherbivores: the influence of very large body size on ecology. Cambridge University Press, Cambridge
Porter WP, Kearney M (2009) Size, shape, and the thermal niche of endotherms. Proc Natl Acad Sci 106:19666–19672. https://doi.org/10.1073/pnas.0907321106
Power JR, Compion SRX (2009) Lion predation on elephants in the Savuti, Chobe national park, Botswana. Afr Zool 44:36–44. https://doi.org/10.1080/15627020.2009.11407437
R Core Team (2020) R: A language and environment for statistical computing. Viena, Austria. Retrieved from https://www.R-project.org/
Ruckstuhl K, Neuhaus P (2000) Sexual segregation in ungulates: a new approach. Behaviour 137:361–377. https://doi.org/10.1163/156853900502123
Ruckstuhl KE, Neuhaus P (2002) Sexual segregation in ungulates: a comparative test of three hypotheses. Biol Rev 77:77–96. https://doi.org/10.1017/s1464793101005814
Ruggiero RG (1991) Opportunistic predation on elephant calves. Afr J Ecol 29:86–89. https://doi.org/10.1111/j.1365-2028.1991.tb00823.x
Shannon G, Page BR, Duffy KJ, Slotow R (2006a) The consequences of body size dimorphism: are African elephants sexually segregated at the habitat scale? Behaviour 143:1145–1168. https://doi.org/10.1163/156853906778607372
Shannon G, Page BR, Duffy KJ, Slotow R (2006b) The role of foraging behaviour in the sexual segregation of the African elephant. Oecologia 150:344–354. https://doi.org/10.1007/s00442-006-0521-1
Shannon G, Page BR, Mackey RL, Duffy KJ, Slotow R (2008) Activity budgets and sexual segregation in African elephants (Loxodonta africana). J Mammal 89:467–476
Shannon G, Thaker M, Vanak AT, Page BR, Grant R, Slotow R (2011) Relative impacts of elephant and fire on large trees in a savanna ecosystem. Ecosystems 14:1372–1381. https://doi.org/10.1007/s10021-011-9485-z
Shrestha A, Van Wieren S, Van Langevelde F, Fuller A, Hetem R, Meyer L, De Bie S, Prins H (2014) Larger antelopes are sensitive to heat stress throughout all seasons but smaller antelopes only during summer in an African semi-arid environment. Int J Biometeorol 58:41–49. https://doi.org/10.1007/s00484-012-0622-y
Sinclair ARE (1995) Serengeti past and present. In: Sinclair ARE, Arcese P (eds) Serengeti II: Dynamics, management, and conservation of an ecosystem. University of Chicago Press, Illinois, USA, pp 3–30
Sinclair ARE, Hopcraft JGC, Olff H, Mduma SAR, Galvin KA, Sharam GJ (2008) Historical and future changes to the Serengeti ecosystem. In: Sinclair ARE, Packer C, Mduma SAR, Fryxell JM (eds) Serengeti III: human impacts on ecosystem dynamics. Chicago University Press, Chicago, pp 7–46
Sinclair ARE, Mduma SAR, Hopcraft JGC, Fryxell JM, Hilborn R, Thirgood S (2007) Long‐term ecosystem dynamics in the Serengeti: lessons for conservation. Conserv Biol 21:580–590. https://doi.org/10.1111/j.1523-1739.2007.00699.x
Singer MS, Bernays EA, Carriere Y (2002) The interplay between nutrient balancing and toxin dilution in foraging by a generalist insect herbivore. Anim Behav 64:629–643. https://doi.org/10.1006/anbe.2002.3082
Skarpe C, du Toit JT, Moe SR (2014) Elephants and savanna woodland ecosystems: a study from Chobe National Park, Botswana. Wiley Blackwell: West Sussex, UK
Skinner JD, Chimimba CT (2005) The mammals of the southern African sub-region. Cambridge University Press, Cambridge
Stokke S (1999) Sex differences in feeding-patch choice in a megaherbivore: elephants in Chobe National Park, Botswana. Can J Zool 77:1723–1732. https://doi.org/10.1139/z99-145
Stokke S, du Toit JT (2000) Sex and size related differences in the dry season feeding patterns of elephants in Chobe National Park, Botswana. Ecography 23:70–80. https://doi.org/10.1111/j.1600-0587.2000.tb00262.x
Stokke S, du Toit JT (2002) Sexual segregation in habitat use by elephants in Chobe National Park, Botswana. Afr J Ecol 40:360–371. https://doi.org/10.1046/j.1365-2028.2002.00395.x
Strauch AM (2011) Seasonal variability in faecal bacteria of semiarid rivers in the Serengeti National Park, Tanzania. Mar Freshw Res 62:1191–1200. https://doi.org/10.1017/s0266467414000625
Strauss MKL, Packer C (2015) Did the elephant and giraffe mediate change in the prevalence of palatable species in an East African Acacia woodland? J Trop Ecol 31:1–12. https://doi.org/10.1017/S0266467414000625
Valls-Fox H (2015) To drink or not to drink? The influence of resource availability on elephant foraging and habitat selection in a semi-arid savanna. (PhD thesis), Université Montpellier, Montpellier, France
Van de Koppel J, Prins HHT (1998) The importance of herbivore interactions for the dynamics of African savanna woodlands: an hypothesis. J Trop Ecol 24:565–576. https://doi.org/10.1017/s0266467498000406
Van Soest PJ (1996) Allometry and ecology of feeding behavior and digestive capacity in herbivores: a review. Zoo Biol 15:455–479. https://doi.org/10.1002/(SICI)1098-2361(1996)15:5%3c455::AID-ZOO3%3e3.0.CO;2-A
Vanak AT, Shannon G, Thaker M, Page B, Grant R, Slotow R (2012) Biocomplexity in large tree mortality: interactions between elephant, fire and landscape in an African savanna. Ecography 35:315–321. https://doi.org/10.1111/j.1600-0587.2011.07213.x
Wittemyer G, Barner Rasmussen H, Douglas-Hamilton I (2007) Breeding phenology in relation to NDVI variability in free-ranging African elephant. Ecography 30:42–50. https://doi.org/10.1111/j.0906-7590.2007.04900.x
Wolanski E, Gereta E (2001) Water quantity and quality as the factors driving the Serengeti ecosystem, Tanzania. Hydrobiologia 458:169–180. https://doi.org/10.1023/A:1013125321838
Woolley LA, Page B, Slotow R (2011) Foraging strategy within African elephant family units: why body size matters. Biotropica 43:489–495. https://doi.org/10.1111/j.1744-7429.2010.00733.x
Woolley LA, Millspaugh JJ, Woods RJ, Van Rensburg SJ, Page BR, Slotow R (2009) Intraspecific strategic responses of African elephants to temporal variation in forage quality. J Wildl Manag 73:827–835. https://doi.org/10.2193/2008-412
Woolnough A, du Toit J (2001) Vertical zonation of browse quality in tree canopies exposed to a size-structured guild of African browsing ungulates. Oecologia 129:585–590. https://doi.org/10.1007/s004420100771
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We thank the Tanzania Wildlife Research Institute (TAWIRI) for offering us a permit to carry this study and the Tanzania National Parks Authority for allowing us to conduct the research in Serengeti National Park. We are thankful to the park staff for cooperating with us during the fieldwork. A permit for conducting the study with a reference number TWRI/RG/22/VOL.39/88/97 was offered by TAWIRI.
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The study was funded by the Norwegian State Education Loan Fund and Inland Norway University of Applied Sciences as a PhD scholarship. The study did not receive any specific grant from funding agencies.
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Mramba, R.P., Mlingi, V.N. The utilisation of woody species by male and female elephants in the Serengeti National Park, a nutrient-rich savanna. Eur J Wildl Res 67, 75 (2021). https://doi.org/10.1007/s10344-021-01523-0
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DOI: https://doi.org/10.1007/s10344-021-01523-0