Predators and management influence adult sex ratio of ungulate communities : synergies and challenges for contemporary conservation
| dc.contributor.author | Abraham, Andrew J. | |
| dc.contributor.author | Panaino, Wendy | |
| dc.contributor.author | Jones, Olivia | |
| dc.contributor.author | Smith, Dylan | |
| dc.contributor.author | Maré, Celesté | |
| dc.contributor.author | Clauss, Marcus | |
| dc.contributor.author | Le Roux, Elizabeth | |
| dc.contributor.author | Webster, Andrea B. | |
| dc.date.accessioned | 2026-04-09T04:43:30Z | |
| dc.date.available | 2026-04-09T04:43:30Z | |
| dc.date.issued | 2025-11 | |
| dc.description | SUPPORTING INFORMATION TABLE S1. Statistical comparison of environmental conditions between the Korannaberg and Lekgaba sections of Tswalu KalahariReserve. Only Mn concentration in trees was found statistically significant (p < 0.05), although this was not considered significant when a Bonferroni correction was applied. TABLE S2. Mean and range [min, max] of abundance and density estimates of large ungulate (>10 kg) species for the period 2020–2023 in the Greater Korannaberg and Lekgaba sections of Tswalu Kalahari Reserve, South Africa. Values in brackets represent minimum and maximum estimates. Information pertaining to sensitive species (rhinos) has been omitted for security reasons. Some species (e.g. buffalo, roan and sable) only occur at low densities in the Greater Korannaberg section and have thus not been included in analysis of adult sex ratios. TABLE S3. Sex bias in predation rate for large African carnivores assembled from our semi-systematic literature review. Sex bias was determined from individual literature sources, where ‘M’ indicates male bias, ‘F’ for female bias and ‘NONE’ for no bias or a preference not statistically significant. * are used to determine results that are statistically significant. Values without a * report a trend in sex bias from qualitative assessments (e.g. reported in paper discussions). FIGURE S1. Environmental comparison between the Korannaberg and Lekgaba sections of Tswalu Kalahari Reserve. (A) Per unit area there are more permanent waterhole sites (and thus artificial mineral lick locations) in Lekgaba. However, per ungulate individual, water/mineral lick availability is remarkably similar. (B) Wilcox-tests reveal no statistically significant (p < 0.05) differences in forage nutrient quality across 10 elements associated with ungulate health, except for Mn in C3 plants (trees). (C) Wilcox-tests also reveal no statistical difference in mean annual rainfall between sections. FIGURE S2. Routes driven during ungulate demographic assessments at Tswalu Kalahari Reserve Black lines display the road network across Tswalu. Routes taken (8 in the Greater Korannaberg section and 4 in the Lekgaba section) are highlighted in red. Effort was made to proportionally cover different habitat types across the reserve. No roads travel over the Korannaberg mountains (habitat: Mtn shrubveld) and thus routes were chosen to travel along the bottom of mountain edges. FIGURE S3. (a) Inter-annual comparison of adult sex ratio (ASR)—represented as percentage of males within each population—of large herbivore species resident within the Korannaberg and Lekgaba sections of Tswalu between 2021 and 2024. Error bars represent 95% confidence intervals generated from 100-fold bootstrapping procedure. Only results where >20 individuals were observed in both sections are reported. For each species, the number of years ASR was significantly different (p < 0.05) based on chi-squared (𝜒2) tests are highlighted. Springbok, impala and blue wildebeest are consistently different across years. Giraffe were not found statistically different in any individual year, although males account for a greater proportion in each field survey and are thus statistically significant when years are combined (see Figure 1). Seasonal comparison of ASR in (b) Korannaberg and (c) Lekgaba for the year 2024, where two field surveys were undertaken at the end of the wet season (February) and end of the dry season (September). In general, results are similar between seasons, although notable discrepancies occur for springbok, impala and eland. We note, however, that these species tend to form large herds segregated by sex. It is therefore possible that observed differences were driven by low sample size rather than actual changes in ungulate ASR throughout the season. FIGURE S4. Comparison of predator–prey preference at Tswalu Kalahari Reserve and continental averages for lion (Panthera leo), cheetah (Acinonyx jubatus) and wild dog (Lycaon pictus). Continental averages taken from Hayward et al. (2005), Hayward et al. (2006a) and Hayward et al. (2006b). FIGURE S5: Sex bias in prey kills for lion, cheetah and wild dog, calculated using Jacob's preference index. Values closer to +1 indicate male bias, while values closer to −1 show female bias. Prey species shown have >20 observations with associated sex information from guide reported predator–prey kill database. FIGURE S6. Seasonal predator consumptive effects on herbivore adult sex ratio at Tswalu. (a) Prey preference of large herbivore species (>10 kg) for lion (Panthera leo), cheetah (Acinonyx jubatus) and African wild dog (Lycaon pictus), calculated using Jacob's preference index by season. Values closer to 1 indicate preferred prey, while values closer to −1 are avoided prey. Summer (wet season) and winter (dry season) seasons are defined by November–April and May–October, respectively. Species are grouped into lion-preferred, all-preferred and cheetah-preferred groups. (b) Seasonal sex bias in prey kills for each prey group by lion, cheetah and wild dog, calculated using Jacob's preference index. Values closer to +1 indicate male bias, while values closer to −1 show female bias. Only groups with n > 10 are shown. Overall, there is remarkably little seasonal variation in either prey preference or sex bias across large carnivore species, except for cheetah preferring female prey during the dry season, but displaying no sex preference during the wet season. FIGURE S7. Predator non-consumptive effects on herbivore adult sex ratio at Tswalu Kalahari Reserve (TKR). (A) Comparison of herbivore nutrition, based on concentration of faecal nitrogen (N), phosphorus (P) and sodium (Na), between sections of TKR for a lion-preferred prey (blue wildebeest; Connochaetes taurinus), all-preferred prey species (kudu; Tragelaphus strepsiceros) and cheetah-preferred species (springbok; Antidorcas marsupialis). (B) Comparison of herbivore body condition between sections of TKR for male and female individuals of large herbivore species. Note: body condition scores have been jittered from 0.5 scoring increments for visual effect. Significant differences between sections for both faecal and body condition scores are denoted, where not significant (ns), p < 0.05 (*), p < 0.01 (**) and p < 0.001 (***). Across both faecal and body condition metrics, predators appear to have little effect on herbivore nutrition and body condition. FIGURE S8. Sex ratio of large herbivores supplemented into the Lekgaba section of Tswalu Kalahari Reserve during the period 2010–2024, compared to the adult sex ratio of large herbivores surveyed in Lekgaba during the period 2021–2024. Numbers to the right highlight the total number of animals supplemented in Lekgaba during 2010–2024. | |
| dc.description | DATA AVAILABILITY STATEMENT : Data available from the Figshare Repository: https://doi.org/10.6084/m9.figsh are.30008 959.v1 (Abraham et al., 2025). | |
| dc.description.abstract | 1. Adult sex ratio (ASR) is important for the ecology, behaviour and life histories of large ungulate populations. ASR, however, varies considerably in wild populations, with driving factors not fully understood. Predation is one possible driver and can alter the ASR of prey species via direct and indirect mechanisms. In the absence of predators, management may also alter ungulate ASR via sex-biased animal removal. 2. Here, we utilised a wildlife reserve divided into two sections, but with comparable environments and ungulate communities. One section hosts lions (Panthera leo) with no management-driven wildlife offtakes, while the other has cheetah (Acinonyx jubatus), wild dogs (Lycaon pictus) and management periodically removes ungulates. To assess the impacts of these differences on ungulate ASR, we surveyed 12 ungulate species (body mass: 10–2000 kg) that co-occur in both sections, revealing that five ungulate species exhibited statistically significant differences in ASR between sections. 3. Predators exerted contrasting direct effects on prey ASR: lion generally displayed a bias towards killing male prey individuals, cheetah had a weak female bias in smaller prey species, and wild dog displayed no general pattern across species. A meta-analysis of sex-biased predator mortality in 53 ungulate populations across Africa broadly supported our findings. Lion appeared to further elicit indirect changes to herding and sexual segregation behaviours in larger-bodied ungulate species, but we found limited evidence for changes in ungulate nutrition or body condition. 4. In the section without lions, wildlife management annually removed ~10%–20% of larger-bodied ungulate populations, with a mild bias towards male individuals. This intervention somewhat replicated the direct effects of natural lion predation on ungulate ASR. Management removals did not, however, stimulate the same behavioural effects in ungulate populations as lions. 5. Synthesis and applications. Predators and management can influence ungulate adult sex ratio (ASR) through multiple pathways, with important implications for breeding system evolution and population dynamics. Given many landscapes are defaunated of natural predators, we demonstrate the importance of incorporating sex-biased wildlife management into contemporary conservation plans. Where management attempts to replicate the effects of ‘missing’ predators, efforts should be made to monitor ungulate ASR and synergistically complement existing predator guilds. | |
| dc.description.department | Mammal Research Institute | |
| dc.description.librarian | am2026 | |
| dc.description.sdg | SDG-15: Life on land | |
| dc.description.sponsorship | H2020 Marie Skłodowska-Curie Actions. | |
| dc.description.uri | https://besjournals.onlinelibrary.wiley.com/journal/13652664 | |
| dc.identifier.citation | Abraham, A. J., Panaino, W., Jones, O., Smith, D., Maré, C., Clauss, M., Le Roux, E., & Webster, A. B. (2025). Predators and management influence adult sex ratio of ungulate communities: Synergies and challenges for contemporary conservation. Journal of Applied Ecology, 62, 3026–3039. https://doi.org/10.1111/1365-2664.70173. | |
| dc.identifier.issn | 0021-8901 (print) | |
| dc.identifier.issn | 1365-2664 (online) | |
| dc.identifier.other | 10.1111/1365-2664.70173 | |
| dc.identifier.uri | http://hdl.handle.net/2263/109479 | |
| dc.language.iso | en | |
| dc.publisher | Wiley | |
| dc.rights | © The Author(s) 2025. This article is licensed under a Creative Commons Attribution 4.0 International License. | |
| dc.subject | Carnivore | |
| dc.subject | Herbivore | |
| dc.subject | Predator | |
| dc.subject | Prey | |
| dc.subject | Prey selection | |
| dc.subject | Sex-bias | |
| dc.subject | Wildlife management | |
| dc.subject | Adult sex ratio (ASR) | |
| dc.title | Predators and management influence adult sex ratio of ungulate communities : synergies and challenges for contemporary conservation | |
| dc.type | Article |
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