Abstract
Common hippopotamus (Hippopotamus amphibius) (hereafter hippos) are iconic and understudied ecological engineers in African aquatic and terrestrial systems. Temporal changes in hippo group size in response to changes in wading areas have been described in river systems, but hippo group dynamics remain generally understudied and have not been quantified in a seasonal floodplain. We used monthly census data collected with an unmanned aerial vehicle between August 2016 and July 2017 to identify changes in hippo distribution, density, and group size at the most used wading areas in the seasonal floodplain in Ndumo Game Reserve, South Africa. We found that hippos congregate into large groups when the wading area decreases in seasonal floodplain systems, similar to perennial river systems. Moreover, as the inundation area increased, overall individual and group density decreased, hippo abundance decreased, and the mean distance between groups increased. Periods of low water availability tend to coincide with periods of food scarcity; thus, competition for water may compound with competition for food, likely affecting cycles of population growth or decline. Increased unpredictability in seasonal and annual rainfall patterns because of climate change will affect the predictability and availability of hippo wading areas, which will likely affect the distribution, size, and persistence of hippo populations. The results of our study provide additional information necessary for the conservation and management of hippos in seasonal floodplain systems in the context of climate change.
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Data for this study are available at: https://doi.org/10.5281/zenodo.4781018. Reproducible code used for the analyses is available at https://doi.org/10.5281/zenodo.6535966.
References
Attwell RIG (1963) Surveying luangwa hippo. Puku 1:29–49
Aureli F, Schaffner CM, Boesch C et al (2008) Fission–fusion dynamics: new research frameworks. Curr Anthropol 49(4):627–654. https://doi.org/10.1086/586708
Bartoń K (2009) MuMIn: multi-model inference. http://r-forge.r-project.org/projects/mumin/
Bercovitch FB, Berry PSM (2013) Herd composition, kinship and fission-fusion social dynamics among wild giraffe. Afr J Ecol 51(2):206–216. https://doi.org/10.1111/aje.12024
Birkhead A, Brown C, Joubert A, Singh A, Tlou T (2018) The Pongola Floodplain, South Africa—part 1: two-dimensional hydrodynamic modelling in support of an environmental flows assessment. Water SA 44:730–745. https://doi.org/10.4314/wsa.v44i4.21
Blackmore A (2017) Legal and public trust considerations for the Ndumo Game Reserve and South Africa-Mozambique border, following the migration of the uSuthu river. S Afr Public Law 30(2):347–378. https://doi.org/10.25159/2522-6800/3585
Brooks ME, Kristensen K, van Benthem KJ, Magnusson A, Berg CW, Nielsen A, Skaug HJ, Maechler M, Bolker BM (2017) glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. R J 9(2):378–400
Chansa W, Milanzi J, Sichone P (2011) Influence of river geomorphologic features on hippopotamus density distribution along the Luangwa River, Zambia: river geomorphology and hippo distribution. Afr J Ecol 49:221–226. https://doi.org/10.1111/j.1365-2028.2010.01252.x
Conradt L, Roper TJ (2000) Activity synchrony and social cohesion: a fission–fusion model. Proc Biol Sci 267:2213–2218. https://doi.org/10.1098/rspb.2000.1271
Dawson J, Pillay D, Roberts PJ, Perissinotto R (2016) Declines in benthic macroinvertebrate community metrics and microphytobenthic biomass in an estuarine lake following enrichment by hippo dung. Sci Rep 6:37359. https://doi.org/10.1038/srep37359
Dawson J, Pillay D, Perissinotto R (2020a) Idiosyncratic responses of meiofaunal assemblages to hippo dung inputs in an estuarine lake. Estuar Coast Shelf Sci 239:106745. https://doi.org/10.1016/j.ecss.2020.106745
Dawson J, Pillay D, Perissinotto R, Richoux NB (2020b) Fatty acid analyses provide novel insights on hippo defecation and consequences for aquatic food webs. Sci Rep 10:12039. https://doi.org/10.1038/s41598-020-68369-5
deJager E (2016) SA rainfall in 2015 the lowest on record. South African Weather Service. https://www.politicsweb.co.za/documents/sa-rainfall-in-2015-the-lowest-on-record--saws. Accessed 4 Nov 2020
Dibloni TO, Vermeulen C, Guenda W, Alfred MN (2010) Structure démographique et mouvements saisonniers des populations d’hippopotame commun, Hippopotamus amphibius Linné 1758 dans la zone sud soudanienne du Burkina faso. Trop Conserv Sci 3:175–189. https://doi.org/10.1177/194008291000300205
Driver A, Sink KJ, Nel JL, Holness S, Van Niekerk L, Daniels F, Jonas Z, Majiedt PA, Harris L, Maze K (2012) National biodiversity assessment 2011: an assessment of South Africa’s biodiversity and ecosystems. Pretoria
Dutton CL, Subalusky AL, Hamilton SK, Rosi EJ, Post DM (2018) Organic matter loading by hippopotami causes subsidy overload resulting in downstream hypoxia and fish kills. Nat Comm 9:1951. https://doi.org/10.1038/s41467-018-04391-6
Dutton CL, Subalusky AL, Hamilton SK, Bayer EC, Njoroge L, Rosi EJ, Post DM (2021) Alternative biogeochemical states of river pools mediated by hippo use and flow variability. Ecosystems 24:284–300. https://doi.org/10.1007/s10021-020-00518-3
Eksteen J, Goodman P, Whyte I, Downs C, Taylor R (2016) A conservation assessment of Hippopotamus amphibius. South African National Biodiversity Institute and Endangered Wildlife Trust, South Africa The Red List of Mammals of South Africa, Swaziland and Lesotho
Field CR (1970) A study of the feeding habits of the hippopotamus (Hippopotamus amphibius Linn.) in the Queen Elizabeth National Park, Uganda, with some management implications. Zool Afr 5:71–86. https://doi.org/10.1080/00445096.1970.11447382
Fouchy K, McClain ME, Conallin J, O’Brien G (2019) Multiple stressors in African freshwater systems. In: Sabater S, Elosegi A, Ludwig R (eds) Multiple stressors in river ecosystems. Elsevier, Hoboken, pp 179–191
Fritsch CJ, Downs CT (2020) Evaluation of low-cost consumer-grade UAVs for conducting comprehensive high-frequency population censuses of hippopotamus populations. Conserv Sci Pract 2:e281. https://doi.org/10.1111/csp2.281
Fritsch CJ, Hanekom C, Downs CT (2021a) Hippopotamus population trends in Ndumo Game Reserve, South Africa, from 1951 to 2021. Glob Ecol Conserv 32:e01910. https://doi.org/10.1016/j.gecco.2021.e01910
Fritsch CJ, Plebani M, Downs CT (2021b) Data from: “Inundation area drives hippo group aggregation and dispersal in a seasonal floodplain system.” Zenodo. https://doi.org/10.5281/zenodo.4781018
Grantham TE, Matthews JH, Bledsoe BP (2019) Shifting currents: managing freshwater systems for ecological resilience in a changing climate. Water Secur 8:100049. https://doi.org/10.1016/j.wasec.2019.100049
Hartig F (2021) DHARMa: residual diagnostics for hierarchical (multi-level/mixed) regression models. R package version 0.4.3. https://CRAN.R-project.org/package=DHARMa
Jacobsen NHG, Kleynhans CJ (1993) The importance of weirs as refugia for hippopotami and crocodiles in the Limpopo River, South Africa. Water SA 19:301–301
Kanga EM, Ogutu JO, Piepho H-P, Olff H (2012) Human–hippo conflicts in Kenya during 1997–2008: vulnerability of a megaherbivore to anthropogenic land use changes. J Land Use Sci 7:395–406. https://doi.org/10.1080/1747423X.2011.590235
Kanga EM, Ogutu JO, Piepho H-P, Olff H (2013) Hippopotamus and livestock grazing: influences on riparian vegetation and facilitation of other herbivores in the Mara Region of Kenya. Landsc Ecol Eng 9:47–58. https://doi.org/10.1007/s11355-011-0175-y
Karstad E, Hudson R (1986) Social organisation and communication of riverine hippopotami in southwestern Kenya. Mammalia 50:153–164
Kelley JL, Morrell LJ, Inskip C, Krause J, Croft DP (2011) Predation risk shapes social networks in fission-fusion populations. PLoS One 6:e24280. https://doi.org/10.1371/journal.pone.0024280
Kilham P (1982) The effect of hippopotamuses on potassium and phosphate ion concentrations in an African lake. Am Midl Nat 108:202. https://doi.org/10.2307/2425311
King J, Brown C, Sabet H (2003) A scenario-based holistic approach to environmental flow assessments for rivers. River Res Appl 19:619–639. https://doi.org/10.1002/rra.709
Klingel H (1979) Social organization of Hippopotamus amphibius. Verhandlungen Dtsch Zool Ges 72:245
Kristensen K, McGillycuddy M (2021) Covariance Structures with GlmmTMB. https://cran.r-project.org/web/packages/glmmTMB/vignettes/covstruct.html. Accessed 21 Sept 2021
Kyle R (1996) Ndumo Game Reserve, South Africa: information sheet for the site designated to the List of Wetlands of International Importance in terms of the Convention on Wetlands of International Importance especially as waterfowl habitat. Pretoria, South Africa: Department of Environmental Affairs and Tourism Report No.: 24/21/3/3/3/15
Lewison RL, Carter J (2004) Exploring behavior of an unusual megaherbivore: a spatially explicit foraging model of the hippopotamus. Ecol Model 171:127–138. https://doi.org/10.1016/S0304-3800(03)00305-3
Lewison R, Pluhacek J (2017) Hippopotamus amphibius. The IUCN red list of threatened species report no.: e. T10103A18567364. https://doi.org/10.2305/IUCN.UK.2017-2.RLTS.T10103A18567364.en
Leyhausen P (1976) Erosion and the hippos. Oryx 13:303–304
Linchant J, Lhoest S, Quevauvillers S, Lejeune P, Vermeulen C, Semeki Ngabinzeke J, Luse Belanganayi B, Delvingt W, Bouché P (2018) UAS imagery reveals new survey opportunities for counting hippos. PLoS One 13:e0206413. https://doi.org/10.1371/journal.pone.0206413
Lock JM (1972) The effects of hippopotamus grazing on grasslands. J Ecol 60:445. https://doi.org/10.2307/2258356
Mackie CS, Dunham KM, Ghiurghi A (2013) Current status and distribution of the vulnerable common hippopotamus Hippopotamus amphibius in Mozambique. Oryx 47:70–76. https://doi.org/10.1017/S0030605311001554
Massey AL, King AA, Foufopoulos J (2014) Fencing protected areas: a long-term assessment of the effects of reserve establishment and fencing on African mammalian diversity. Biol Conserv 176:162–171. https://doi.org/10.1016/j.biocon.2014.05.023
McCarthy TS, Ellery WN, Bloem A (1998) Some observations on the geomorphological impact of hippopotamus (Hippopotamus amphibius L.) in the Okavango Delta. Botswana Afr J Ecol 36:44–56
McCauley DJ, Dawson TE, Power ME et al (2015) Carbon stable isotopes suggest that hippopotamus-vectored nutrients subsidise aquatic consumers in an East African river. Ecosphere 6:art52. https://doi.org/10.1890/ES14-00514.1
McCauley DJ, Graham SI, Dawson TE, Power ME, Ogada M, Nyingi WD, Githaiga JM, Nyunja J, Hughey LF, Brashares JS (2018) Diverse effects of the common hippopotamus on plant communities and soil chemistry. Oecologia 188:821–835. https://doi.org/10.1007/s00442-018-4243-y
Nakagawa S, Schielzeth H (2013) A general and simple method for obtaining R2 from generalized linear mixed-effects models. Methods Ecol Evol 4:133–142
O’Brien GC, Ross M, Hanzen C, Dlamini V, Petersen R, Diedericks GJ, Burnett MJ (2019) River connectivity and fish migration considerations in the management of multiple stressors in South Africa. Mar Freshw Res 70:1254. https://doi.org/10.1071/MF19183
O’Connor TG, Campbell BM (1986) Hippopotamus habitat relationships on the Lundi River, Gonarezhou National Park, Zimbabwe. Afr J Ecol 24:7–26
Olivier RCD, Laurie WA (1974) Habitat utilization by hippopotamus in the Mara River. Afr J Ecol 12:249–271. https://doi.org/10.1111/j.1365-2028.1974.tb01036.x
Onditi KO, Li X, Song W, Li Q, Musila S, Mathenge J, Kioko E, Jiang X (2021) The management effectiveness of protected areas in Kenya. Biodivers Conserv 30:3813–3836. https://doi.org/10.1007/s10531-021-02276-7
Pekor A, Miller JR, Flyman MV, Kasiki S, Kesch MK, Miller SM, Uiseb K, Van der Merve V, Lindsey PA (2019) Fencing Africa’s protected areas: Costs, benefits, and management issues. Biol Conserv 229:67–75. https://doi.org/10.1016/j.biocon.2018.10.030
Plebani M, Fritsch CJ, Downs CT (2022) Reproducible statistical analyses for the article: "Inundation area drives hippo group aggregation and dispersal in a seasonal floodplain system". Zenodo. https://doi.org/10.5281/zenodo.6535966
Pooley S (2021) Biodiversity hotspot and Ramsar site under threat. Oryx 55:811–812. https://doi.org/10.1017/S0030605321001174
Prinsloo AS, Pillay D, O’Riain MJ (2020) Multiscale drivers of hippopotamus distribution in the St Lucia Estuary, South Africa. Afr Zool 55:127–140. https://doi.org/10.1080/15627020.2020.1717377
R Core Team (2021) R: a language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. https://www.R-project.org/
Ramos-Fernández G, Morales JM (2014) Unraveling fission-fusion dynamics: how subgroup properties and dyadic interactions influence individual decisions. Behav Ecol Sociobiol 68:1225–1235. https://doi.org/10.1007/s00265-014-1733-8
Schoelynck J, Subalusky AL, Struyf E, Dutton CL, Unzué-Belmonte D, Van de Vijver B, Post DM, Rosi EJ, Meire P, Frings P (2019) Hippos (Hippopotamus amphibius): the animal silicon pump. Sci Adv 5:aav0395. https://doi.org/10.1126/sciadv.aav0395
Scholte P, Nguimkeng F, Iyah E (2017) Good news from north-central Africa: largest population of vulnerable common hippopotamus Hippopotamus amphibius is stable. Oryx 51:218–221. https://doi.org/10.1017/S0030605315001258
Sidney J (1965) The past and present distribution of some African ungulates. Trans Zool Soc Lond 30:1–397
Smit IPJ, Archibald S (2019) Herbivore culling influences spatio-temporal patterns of fire in a semiarid savanna. J Appl Ecol 56:711–721. https://doi.org/10.1111/1365-2664.13312
Smit IPJ, Peel MJ, Ferreira SM, Greaver C, Pienaar DJ (2020) Megaherbivore response to droughts under different management regimes: lessons from a large African savanna. Afr J Range Forage Sci 37:65–80. https://doi.org/10.2989/10220119.2019.1700161
Smuts GL, Whyte IJ (1981) Relationships between reproduction and environment in the hippopotamus Hippopotamus amphibius in the Kruger National Park. Koedoe 24:169–185
Stears K, McCauley DJ, Finlay JC, Mpemba J, Warrington IT, Mutayoba BM, Power ME, Dawson TE, Brashares JS (2018) Effects of the hippopotamus on the chemistry and ecology of a changing watershed. Proc Natl Acad Sci 115:E5028–E5037. https://doi.org/10.1073/pnas.1800407115
Stears K, Nuñez TA, Muse EA, Mutayoba BM, McCauley DJ (2019) Spatial ecology of male hippopotamus in a changing watershed. Sci Rep 9:15392. https://doi.org/10.1038/s41598-019-51845-y
Stommel C, Hofer H, East ML (2016) The effect of reduced water availability in the Great Ruaha River on the vulnerable common hippopotamus in the Ruaha National Park, Tanzania. PLoS One 11:e0157145. https://doi.org/10.1371/journal.pone.0157145
Subalusky AL, Dutton CL, Rosi-Marshall EJ, Post DM (2014) The hippopotamus conveyor belt: vectors of carbon and nutrients from terrestrial grasslands to aquatic systems in sub-Saharan Africa. Freshw Biol 60:512–525. https://doi.org/10.1111/fwb.12474
Subalusky AL, Anderson EP, Jiménez G et al (2019) Potential ecological and socio-economic effects of a novel megaherbivore introduction: the hippopotamus in Colombia. Oryx 55:105–113. https://doi.org/10.1017/S0030605318001588
Tockner K, Stanford JA (2002) Review of riverine flood plains: present state and future trends. Env Con 24:308–330
Verweij RJT, Verrelst J, Loth PE, Heitkönig IMA, Brunsting AMH (2006) Grazing lawns contribute to the subsistence of mesoherbivores on dystrophic savannas. Oikos 114:108–116
Viljoen PC (1980) Distribution and numbers of hippopotamus in the Olifants and Blyde Rivers. South Afr J Wildl Res 10:129–132
Viljoen PC (1995) Changes in number and distribution of hippoptamus (Hippopotamus amphibius) in the Sabie River, Kruger National Park, during the 1992 drought. Koedoe 38:115–121
Viljoen PC, Biggs HC (1998) Population trends of hippopotami in the rivers of the Kruger National Park, South Africa. In: Dunstone N, Gorman ML (Eds) Behaviour and ecology of riparian mammals, vol 71. Symposia of the Zoological Society of LondonCambridge University Press, . Cambridge, p 251–279. https://doi.org/10.1017/CBO9780511721830.016.
Wielgus E, Cornélis D, de Garine-Wichatitsky M, Cain B, Fritz H, Miguel E, Valls-Fox H, Caron A, Chamaillé-Jammes S (2020) Are fission–fusion dynamics consistent among populations? A large-scale study with Cape buffalo. Ecol Evol 10:9240–9256. https://doi.org/10.1002/ece3.6608
Zisadza P, Gandiwa E, van der Westhuizen H, van der Westhuizen E, Bodzo V (2010) Abundance, distribution and population trends of hippopotamus in Gonarezhou National Park, Zimbabwe. South Afr J Wildl Res 40:149–157. https://doi.org/10.3957/056.040.0206
Zoeller K, Bond WJ (2013) Hippos as ecosystem engineers? Grazing lawns and their determinants in the St Lucia floodplain. S Afr J Bot 86:144
Zuur A, Ieno EN, Walker N, Saveliev AA, Smith GM (2009) Mixed effects models and extensions in ecology with R. Statistics for biology and health. Springer, New York. https://doi.org/10.1007/978-0-387-87458-6
Acknowledgements
We are grateful to Ezemvelo KZN Wildlife, C. Hanekom, and A. Thembe for permission to work in NGR. We would also like to thank all the rangers at NGR and especially B. Nyathi. In addition, this work would not have been possible without the help of all the research assistants involved, in particular C. Price, S. McPherson, V. Thabethe, H. Aronsson, T. Caine, and H. Thatcher. We thank the National Research Foundation (ZA, Grant 98404), the University of KwaZulu-Natal (ZA), the Rufford Trust (UK), the Phoenix Zoo (USA) and the Hans Hoheisen Charitable Trust (ZA) for funding. We thank the Ford Wildlife Foundation (ZA) for vehicle support. MP was supported by grants from the Claude Leon Foundation and the Swiss National Science Foundation. We thank M. Thornhill for his UAV expertise and for servicing the UAV. We thank Planet Imagery for supplying satellite imagery. We thank the reviewers for their constructive comments that improved the manuscript.
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This work was supported by National Research Foundation (ZA, Grant 98404), the Rufford Trust (UK, 0e95d7-1), the Phoenix Zoo (USA), the Hans Hoheisen Charitable Trust (ZA), and the Ford Wildlife Foundation (ZA). MP was funded by the Claude Leon Foundation (ZA) and the Swiss National Science Foundation (CH).
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Fritsch, C.J.A., Plebani, M. & Downs, C.T. Inundation area drives hippo group aggregation and dispersal in a seasonal floodplain system. Mamm Biol 102, 1811–1821 (2022). https://doi.org/10.1007/s42991-022-00286-8
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DOI: https://doi.org/10.1007/s42991-022-00286-8