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Edited by James A. Estes, University of California, Santa Cruz, CA, and approved May 16, 2017 (received for review September 8, 2016)
Article Figures & SI
Figures
- Fig. 1.
Map showing the Mara River Basin, Kenya, and its relationship to the Maasai Mara National Reserve (MMNR) and Serengeti National Park (SNP) (A), the distribution of wildebeest crossing sites along the Mara River coded by the number of mass drownings confirmed to have occurred at each site (for 21 of the 23 drownings) (B), and the distribution of 3,380 wildebeest carcasses over 5 km of river after a mass drowning in July 2011 at the crossing site coded red in B (C).
- Fig. 2.
Input and fate of wildebeest carcasses from mass drownings in the Mara River. (A) Percentage of dry mass composition of an average wildebeest carcass, with stoichiometric ratios by mass shown as C/N/P. (B) Microbial decomposition rates for wildebeest bone, skin, intestines, and muscle in the Mara River (mean ± SEM). (C) Daily estimations of total nitrogen (TN) vs. total phosphorus (TP) flux downstream of three carcass aggregations ranging from 1,000–3,400 carcasses. (D) Declines in NH4+-N concentration downstream of wildebeest carcasses and estimations of nutrient uptake length (Sw) on days 8, 16, and 26 after the drowning. (E) Total dry mass of wildebeest carcasses consumed by avian scavengers on aggregations of 16 and 40 carcasses. (F) Assimilation of wildebeest tissue (in November 2013, 1 mo after carcasses were present) and bone biofilm (in February 2014, 4 mo after carcasses were present) by three common fish taxa (mean ± 95% credibility intervals).
- Fig. 3.
Illustrative diagram showing estimated annual loading and transport pathways of C, N, and P from mass drownings of wildebeest in the Mara River, Kenya. Mean ± SD total loading estimations are at the top of the diagram in terms of carcasses and metric tons of C, N, and P, and the range of estimations (minimum–maximum) for the amount transported in each pathway are presented as the percentage of total loading. A through F align with data presented in Fig. 2. Fluxes through the two pathways illustrated in F are currently unknown.
Tables
- Table S1.
Number and elemental composition of wildebeest carcasses entering the Mara River from 2011 to 2015
Year No. of mass drownings No. of carcasses Carcass dry mass,* 103 kg Average aerial loading,† g of DM per m−2 C, 103 kg N, 103 kg P, 103 kg 2011 3 8,000 385 590 137 32 16 2012 6 9,400 453 250 161 37 19 2013 7 7,750 373 360 133 31 16 2014 4 2,700 130 640 46 11 5 2015 3 3,400 164 140 58 13 7 Mean (SD) 4.6 (1.8) 6,250 (3,000) 301 (144) 400 (210) 107 (51) 25(12) 13(6) Carcass element Percentage of carcass,* mean (SD) C/N/P† Decay rate, k⋅d−1 (95% CI) Days to 95% biomass loss, mean days (95% CI) Stomach contents 12.4 (2.9) 69.2:3.3:1.0 Muscle 25.7 (0.7) 152.0:45.7:1.0 −0.188 (−0.168 to −0.208) 16 (14–18) Internal organs‡ 7.2 (1.7) 96.4:21.2:1.0 −0.068 (−0.047 to −0.089) 44 (38–63) Skin 10.9 (2.0) 215.5:72.5:1.0 −0.043 (−0.038 to −0.048) 70 (63–80) Bone 43.7 (3.7) 2.6:0.5:1.0 −0.001 (−0.0013 to −0.0009) 2,709 (2,285–3,327) Total carcass 100.0 8.5:2.0:1.0 Date No. of carcasses Average Q, m3⋅s−1 Upstream DOC flux, kg Total carcass C loading, kg Soft tissue C loading, kg Net DOC flux, kg Upstream TN flux, kg Total carcass N loading, kg Soft tissue N loading, kg Net TN flux, kg Upstream TP flux, kg Total carcass P loading, kg Soft tissue P loading, kg Net TP flux, kg 7/4/2011 3,380 9.7 30,310 57,900 40,800 −1,290 17,080 13,400 10,000 1,270 1,510 6,800 320 50 11/5/2012 990 15.6 96,270 17,000 12,000 3,480 64,290 3,900 2,900 −180 6,450 2,000 90 −40 9/25/2013 1,610 16.6 10,8490 27,600 19,400 8,350 60,720 6,400 4,700 2,240 6,250 3,300 150 −210 Mean (SD) 1,990 (1,240) 13.9 (3.7) 78,360 (42,060) 34,200 (21,300) 24,100 (15,000) 3,510 (4,820) 47,370 (26,290) 7,900 (4,900) 5,900 (3,700) 1,110 (1,220) 4,740 (2,790) 4,000 (2,500) 190 (120) −65 All fluxes were calculated over a period of 25–29 d. Q, discharge; TN, total nitrogen.
Species Average mass, kg Daily energy expenditure, kJ⋅d−1 Daily energy consumed, kJ⋅d−1 Daily dry mass consumed, g⋅d−1 Daily C consumed, g⋅d−1 Daily N consumed, g⋅d−1 Daily P consumed, g⋅d−1 Maribou stork 6.45 2,577 2,990 115 51 13 0.4 White-backed vulture 5.43 2,321 2,692 104 46 11 0.4 Rüppell's vulture 7.40 2,803 3,251 125 56 14 0.4 Hooded vulture 2.04 1,277 14,812 57 25 6 0.2
Data supplements
Supporting Information
Wildebeest mass drownings alter river ecosystem
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