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Trophic rewilding revives biotic resistance to shrub invasion


Trophic rewilding seeks to rehabilitate degraded ecosystems by repopulating them with large animals, thereby re-establishing strong top-down interactions. Yet there are very few tests of whether such initiatives can restore ecosystem structure and functions, and on what timescales. Here we show that war-induced collapse of large-mammal populations in Mozambique’s Gorongosa National Park exacerbated woody encroachment by the invasive shrub Mimosa pigra—considered one of the world’s 100 worst invasive species—and that one decade of concerted trophic rewilding restored this invasion to pre-war baseline levels. Mimosa occurrence increased between 1972 and 2015, a period encompassing the near extirpation of large herbivores during the Mozambican Civil War. From 2015 to 2019, mimosa abundance declined as ungulate biomass recovered. DNA metabarcoding revealed that ruminant herbivores fed heavily on mimosa, and experimental exclosures confirmed the causal role of mammalian herbivory in containing shrub encroachment. Our results provide mechanistic evidence that trophic rewilding has rapidly revived a key ecosystem function (biotic resistance to a notorious woody invader), underscoring the potential for restoring ecological health in degraded protected areas.

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Fig. 1: Study system, experimental design, and hypotheses.
Fig. 2: Trends in ungulate biomass, M. pigra abundance, and rainfall.
Fig. 3: Consumption of M. pigra by LMH in the Urema floodplain.
Fig. 4: Large herbivores suppressed vital rates of M. pigra.
Fig. 5: Large herbivores regulate the density and biomass of M. pigra.

Data availability

The field data are provided in Supplementary Data 18. The field data along with raw dietary sequence data and metadata from 2013, 2015, 2017, and 2018 are available via Dryad ( Dietary sequence data and metadata from 2016, along with the local plant reference database, are available via Dryad (


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We thank Parque Nacional da Gorongosa and the government of Mozambique for permission to conduct this research. We thank the Gorongosa Project for facilitating scientific research, with special thanks to M. Marchington, F. Moniz, A. Dos Santos, T. Massad, and G. Carr. The Gorongosa Project had no role in the conceptualization, design, data collection, data analysis, manuscript preparation, or decision to publish. We are indebted to K. Tinley for his pioneering research38 on Gorongosa’s pre-war ecology. Supplementary Video 1 was produced in collaboration with National Geographic Labs Crittercam. Funding was provided by National Geographic Young Explorers Grant no. 9459-14; the US National Science Foundation (grant no. IOS-1656527 and the Graduate Research Fellowship Program); the Princeton Environmental Institute’s Grand Challenges programme; the Randall and Mary Hack ’69 Award for Water and the Environment; Princeton University’s Institutes for African Studies and International and Regional Studies; the Greg Carr Foundation; the Cameron Schrier Foundation; the Sherwood Foundation; and Princeton’s Innovation Fund for New Ideas in the Natural Sciences.

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J.A.G. and R.M.P. conceived and designed the research. J.A.G. conducted the primary plant surveys and the exclosure experiment, and collected specimens for the plant reference database. J.P., M.C.H., and T.R.K. conducted the DNA metabarcoding analyses. J.P. analysed the dietary data. A.B.P. contributed the forage-quality data. M.J.S.P. contributed additional plant-survey data. M.E.S. contributed the wildlife-count data. J.A.G., J.P., M.C.H., T.R.K., T.C.C., J.H.D., A.G.C., and R.M.P. collected samples and field data. J.A.G. and R.M.P. analysed the field data. J.A.G., J.P., and R.M.P. wrote the manuscript. All authors made revisions and approved the final draft.

Corresponding author

Correspondence to Robert M. Pringle.

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Competing interests

A.G.C. and M.E.S. were employed by the Gorongosa Project, a non-profit organization that co-manages conservation and restoration in Gorongosa National Park in partnership with the government of Mozambique. M.J.S.P. was contracted by the Gorongosa Project to conduct vegetation surveys. R.M.P. was an unpaid member on the board of directors of the Gorongosa Project. All other authors have no competing interests.

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Extended data

Extended Data Fig. 1 Top food plants for six dominant ungulates in the Urema floodplain between 2013 and 2018.

Bars show the mean relative read abundance of each plant taxon across all fecal samples in each year for each species. Sample sizes for each species in each year are shown in Fig. 3a. The best possible taxonomic identification for each plant (see Methods) is provided beneath each bar, and the corresponding plant life-form (grass, shrub, tree, or forb) is listed above each bar. Stars within bars denote Mimosa pigra (the first or second most abundant food for all ruminant species in all years).

Extended Data Fig. 2 Mimosa consumption in early (June–August) versus late (October–November) dry season of 2017 (below-average rainfall year) and 2018 (above-average rainfall year).

Bars show the mean (±1 s.e.m.) relative read abundance of Mimosa pigra across all fecal samples in each season for each species. Sample sizes for each species in each season are shown above bars. As for Fig. 3, quantitative comparisons between years (and between seasons for 2018) should be interpreted cautiously (see Methods). Although sample sizes are limited for some species in some seasons, the data show that antelope species consumed mimosa in appreciable quantities throughout the dry season.

Extended Data Fig. 3 Suppression of Mimosa pigra reproduction by large herbivores.

Here, the results in Fig. 4c, d are broken down to show independent trends in immature floral buds (a) and mature flowers (b), along with immature green fruits (c) and mature brown seed pods (d) in the experimental exclosure and control plots. Points show the mean (±1 s.e.m.) number of reproductive structures per plant in each treatment over three years. As in Fig. 4a–d, measurements at the level of individual plants were averaged at the plot level before the analysis (from left to right in each panel, n = 12, 6, 12, 9, 10, and 9 plots per survey). These data show that large herbivores have essentially eliminated reproductive output by mimosa in Gorongosa: few reproductive structures at any stage of development were recorded in the control plots as of 2017, and none at all were found in 2018.

Extended Data Fig. 4 Estimation of aboveground dry biomass from field measurements of plant volume.

Plant volume was calculated using measurements of height and canopy dimensions for each of 34 Mimosa pigra individuals, assuming an ellipsoidal shrub shape, and regressed against the dry aboveground biomass measured for each of the same plants (see Methods). The regression equation shown was used to estimate the aboveground biomass of standing plants in each experimental exclosure and control plot in 2018 (see Fig. 5b).

Extended Data Fig. 5 Rapid recruitment and growth of Mimosa pigra inside, but not outside, experimental herbivore exclosures.

All photos are from the same exclosure-control pair (in long-term monitoring plot 16). a, Panoramic photograph of the control plot in 2018, showing floodplain dominated by grasses (mostly Cynodon dactylon) and forbs (mostly Heliotropium spp.); a total of 13 mimosa plants were recorded in this 260-m2 plot in 2018, none taller than 43 cm. The exclosure plot is visible at top center. b, Forb-dominated understory in the exclosure plot in September 2017, when a total of 57 small mimosa plants were recorded, none taller than 31 cm (up from just one individual recorded in September 2016). ce, Three views of the same exclosure plot in August 2018, when 661 mimosa plants of at least 15-cm stem length were recorded, including individuals up to 158-cm tall.

Supplementary information

Reporting Summary

Supplementary Data 1–8

Field data.

Supplementary Video 1

Waterbuck foraging on Mimosa pigra in the Urema floodplain (8 August 2015). This footage was obtained from a National Geographic Labs Crittercam fitted around the animal’s neck. Note that this individual is foraging within one of the many mudflats that occur at the edges of Lake Urema and adjoining ponds and drainage channels (specific location 18°52’43.82”S, 34°27’26.55”E); these mudflats typically have sparse, forb-dominated plant communities. The limited grass cover evident in the video should not be interpreted as evidence that grass or other forages were limited in the floodplain at large.

Supplementary Video 2

Oribi foraging on Mimosa pigra in the Urema floodplain (23 July 2019).

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Guyton, J.A., Pansu, J., Hutchinson, M.C. et al. Trophic rewilding revives biotic resistance to shrub invasion. Nat Ecol Evol 4, 712–724 (2020).

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