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Flux across the carbon cycle is generally characterized by contributions from plants, microbes, and abiotic systems. Animals, however, move vast amounts of carbon, both through ecosystem webs and across the landscape. Schmitz et al. review the different contributions that animal populations make to carbon cycling and discuss approaches that allow for better monitoring of these contributions. Science , this issue p. eaar3213 Predicting and managing the global carbon cycle requires scientific understanding of ecosystem processes that control carbon uptake and storage. It is generally assumed that carbon cycling is sufficiently characterized in terms of uptake and exchange between ecosystem plant and soil pools and the atmosphere. We show that animals also play an important role by mediating carbon exchange between ecosystems and the atmosphere, at times turning ecosystem carbon sources into sinks, or vice versa. Animals also move across landscapes, creating a dynamism that shapes landscape-scale variation in carbon exchange and storage. Predicting and measuring carbon cycling under such dynamism is an important scientific challenge. We explain how to link analyses of spatial ecosystem functioning, animal movement, and remote sensing of animal habitats with carbon dynamics across landscapes.

Predator-prey relationships are integral to ecosystem stability and functioning. These relationships are, however, difficult to maintain in protected areas where large predators are increasingly being reintroduced and confined. Where predators make kills has a profound influence on their role in ecosystems, but the relative importance of environmental variables in determining kill sites, and how these might vary across ecosystems is poorly known. We investigated kill sites for lions in South Africa's thicket biome, testing the importance of vegetation structure for kill site locations compared to other environmental variables. Kill sites were located over four years using GPS telemetry and compared to non-kill sites that had been occupied by lions, as well as to random sites within lion ranges. Measurements of 3D vegetation structure obtained from Light Detection and Ranging (LiDAR) were used to calculate the visible area (viewshed) around each site and, along with wind and moonlight data, used to compare kill sites between lion sexes, prey species and prey sexes. Viewshed area was the most important predictor of kill sites (sites in dense vegetation were twice as likely to be kill sites compared to open areas), followed by wind speed and, less so, moonlight. Kill sites for different prey species varied with vegetation structure, and male prey were killed when wind speeds were higher compared to female prey of the same species. Our results demonstrate that vegetation structure is an important component of predator-prey interactions, with varying effects across ecosystems. Such differences require consideration in terms of the ecological roles performed by predators, and in predator and prey conservation.

The flexibility of UAV‐lidar remote sensing offers a myriad of new opportunities for savanna ecology, enabling researchers to measure vegetation structure at a variety of temporal and spatial scales. However, this flexibility also increases the number of customizable variables, such as flight altitude, pattern, and sensor parameters, that, when adjusted, can impact data quality as well as the applicability of a dataset to a specific research interest. To better understand the impacts that UAV flight patterns and sensor parameters have on vegetation metrics, we compared 7 lidar point clouds collected with a Riegl VUX − 1LR over a 300 × 300 m area in the Kruger National Park, South Africa. We varied the altitude (60 m above ground, 100 m, 180 m, and 300 m) and sampling pattern (slowing the flight speed, increasing the overlap between flightlines and flying a crosshatch pattern), and compared a variety of vertical vegetation metrics related to height and fractional cover. Comparing vegetation metrics from acquisitions with different flight patterns and sensor parameters, we found that both flight altitude and pattern had significant impacts on derived structure metrics, with variation in altitude causing the largest impacts. Flying higher resulted in lower point cloud heights, leading to a consistent downward trend in percentile height metrics and fractional cover. The magnitude and direction of these trends also varied depending on the vegetation type sampled (trees, shrubs or grasses), showing that the structure and composition of savanna vegetation can interact with the lidar signal and alter derived metrics. While there were statistically significant differences in metrics among acquisitions, the average differences were often on the order of a few centimetres or less, which shows great promise for future comparison studies. We discuss how these results apply in practice, explaining the potential trade‐offs of flying at higher altitudes and with alternate patterns. We highlight how flight and sensor parameters can be geared toward specific ecological applications and vegetation types, and we explore future opportunities for optimizing UAV‐lidar sampling designs in savannas. A cross‐section of a lidar point cloud for a single tree (left) from a savanna in the Satara region of Kruger National Park, South Africa (visualised with CloudCompare 2.11). Smoothed vertical profiles of fractional canopy cover per 10 cm height bin are plotted for the same tree. These profiles were derived from a series of airborne lidar data collected from 4 different flight altitudes (60 m, 100 m, 180 m, and 300 m above ground) with an unoccupied aerial vehicle (UAV). As flight altitude increases (left‐right), the canopy cover profiles change shape and shift downward, demonstrating that UAV fight and sensor parameters can have a significant impact on lidar measurements of vegetation structure.

The conservation of charismatic and functionally important large species is becoming increasingly difficult. Anthropogenic pressures continue to squeeze available habitat and force animals into degraded and disturbed areas. Ensuring the long-term survival of these species requires a well-developed understanding of how animals use these new landscapes to inform conservation and habitat restoration efforts. We combined 3 y of highly detailed visual observations of Bornean orangutans with high-resolution airborne remote sensing (Light Detection and Ranging) to understand orangutan movement in disturbed and fragmented forests of Malaysian Borneo. Structural attributes of the upper forest canopy were the dominant determinant of orangutan movement among all age and sex classes, with orangutans more likely to move in directions of increased canopy closure, tall trees, and uniform height, as well as avoiding canopy gaps and moving toward emergent crowns. In contrast, canopy vertical complexity (canopy layering and shape) did not affect movement. Our results suggest that although orangutans do make use of disturbed forest, they select certain canopy attributes within these forests, indicating that not all disturbed or degraded forest is of equal value for the long-term sustainability of orangutan populations. Although the value of disturbed habitats needs to be recognized in conservation plans for wide-ranging, large-bodied species, minimal ecological requirements within these habitats also need to be understood and considered if long-term population viability is to be realized.

Tropical savannas have been increasingly targeted for carbon sequestration by afforestation, assuming large gains in soil organic carbon (SOC) with increasing tree cover. Because savanna SOC is also derived from grasses, this assumption may not reflect real changes in SOC under afforestation. However, the exact contribution of grasses to SOC and the changes in SOC with increasing tree cover remain poorly understood. Here we combine a case study from Kruger National Park, South Africa, with data synthesized from tropical savannas globally to show that grass-derived carbon constitutes more than half of total SOC to a soil depth of 1 m, even in soils directly under trees. The largest SOC concentrations were associated with the largest grass contributions (>70% of total SOC). Across the tropics, SOC concentration was not explained by tree cover. Both SOC gain and loss were observed following increasing tree cover, and on average SOC storage within a 1-m profile only increased by 6% (s.e. = 4%, n = 44). These results underscore the substantial contribution of grasses to SOC and the considerable uncertainty in SOC responses to increasing tree cover across tropical savannas.Grasses contribute more than half of the soil organic carbon across tropical savannas, according to a case study in South Africa combined with a synthesis of data from tropical savannas globally.

Humans pose a major threat to many species through land-use change in virtually every habitat. However, the extent of this threat is largely unknown for invertebrates due to challenges with investigating their distributions at large scales. This knowledge gap is particularly troublesome for soil macrofauna because of the critical roles many of these organisms perform as ecosystem engineers. We used a combination of high-resolution airborne Light Detection and Ranging and deep learning models to map the distribution of the ecologically important termite genus Macrotermes across a South African savanna land-use gradient, quantifying the effects of land-use change on patterns of mound densities, heights and spatial patterning. Despite significant anthropogenic alteration to landscapes, termite mounds persisted and shared a number of similarities to mounds in untransformed areas. Mean mound height was not substantially reduced in transformed landscapes, and over-dispersion of mounds at localized scales was conserved. However, mound densities were partially reduced, and height distributions in transformed areas differed to those in protected areas. Our findings suggest that mound-building termites persist even in areas of relatively high human disturbance, but also highlight important differences in termite distributions that could lead to reductions in ecosystem services provided by termites in human-modified landscapes. The persistence of at least half of mounds in human-modified landscapes could serve as starting points for savanna restoration.

Termite mounds contribute to the spatial heterogeneity of ecological processes in many savannas, but the underlying patterns and determinants of mound distributions remain poorly understood. Using the Carnegie Airborne Observatory (CAO), we mapped the distribution of termite mounds across a rainfall gradient within a river catchment (∼ 27 000 ha) of the Kruger National Park, South Africa. We assessed how different factors were associated with the distribution and height of termite mounds at three spatial scales: the entire catchment, among three broad vegetation types, and on individual hillslope crests. Abiotic factors such as the underlying geology and mean annual precipitation shaped mound densities at broad scales, while local hillslope morphology strongly influenced mound distribution at finer scales, emphasising the importance of spatial scale when assessing mound densities. Fire return period had no apparent association with mound densities or height. Mound density averaged 0.46 mounds ha⁻¹, and exhibited a clustered pattern throughout the landscape, occurring at relatively high densities (up to 2 mounds ha⁻¹) on crests, which are nutrient‐poor elements of the landscape. Mounds exhibited significant over‐dispersion (even spacing) at scales below 60 m so that evenly spaced aggregations of termite mounds are embedded within a landscape of varying mound densities. The tallest mounds were found in dry savanna (500 mm yr⁻¹) and were positively correlated with mound density, suggesting that dry granitic savannas are ideal habitat for mound‐building termites. Mound activity status also varied significantly across the rainfall gradient, with a higher proportion of active (live) mounds in the drier sites. The differential spacing of mounds across landscapes provides essential nutrient hotspots in crest locations, potentially sustaining species that would otherwise not persist. The contribution to biodiversity and ecosystem functioning that mounds provide is not uniform throughout landscapes, but varies considerably with spatial scale and context.

Riparian ecosystems are amongst the most biodiverse tropical habitats. They are important, and essential, ecological corridors, linking remnant forest fragments. In this study, we hypothesised that crocodile's actively select nocturnal resting locations based on increased macaque predation potential. We examined the importance of riparian vegetation structure in the maintenance of crocodile hunting behaviours. Using airborne Light Detection and Ranging (LiDAR) and GPS telemetry on animal movement, we identified the repeated use of nocturnal resting sites by adult estuarine crocodiles (Crocodylus porosus) throughout the fragmented Lower Kinabatangan Wildlife Sanctuary in Sabah, Malaysia. Crocodile resting locations were found to resemble, in terms of habitat characteristics, the sleeping sites of long-tailed macaque; positioned in an attempt to avoid predation by terrestrial predators. We found individual crocodiles were actively selecting overhanging vegetation and that the protrusion of trees from the tree line was key to site selection by crocodiles, as well as influencing both the presence and group size of sleeping macaques. Although these findings are correlational, they have broad management implications, with the suggestion that riparian corridor maintenance and quality can have implications beyond that of terrestrial fauna. We further place our findings in the context of the wider ecosystem and the maintenance of trophic interactions, and discuss how future habitat management has the potential to mitigate human-wildlife conflict.

Herbivores do not forage uniformly across landscapes, but select for patches of higher nutrition and lower predation risk. Macrotermes mounds contain higher concentrations of soil nutrients and support grasses of higher nutritional value than the surrounding savanna matrix, attracting mammalian grazers that preferentially forage on termite mound vegetation. However, little is known about the spatial extent of such termite influence on grazing patterns and how it might differ in time and space. We measured grazing intensity in three African savanna types differing in rainfall and foliar nutrients and predicted that the functional importance of mounds for grazing herbivores would increase as the difference in foliar nutrient levels between mound and savanna matrix grasses increases and the mounds become more attractive. We expected this to occur in nutrient‐poor areas and during the dry season when savanna matrix grass nutrient levels are lower. Tuft use and grass N and P content were measured along transects away from termite mounds, enabling calculation of the spatial extent of termite influence on mammalian grazing. Using termite mound densities estimated from airborne light detection and ranging (LiDAR), we further upscaled field‐based results to determine the percentage of the landscape influenced by termite activity. Grasses in close proximity to termite mounds were preferentially grazed at all sites and in both seasons, but the strength of mound influence varied between savanna types and seasons. In the wet season, mounds had a relatively larger effect on grazers at the landscape scale in the nutrient‐poor, wetter savanna, whereas in the dry season the pattern was reversed with more of the landscape influenced at the nutrient‐rich, driest site. Our results reveal that termite mounds enhance the value of savanna landscapes for herbivores, but that their functional importance varies across savanna types and seasons.

Termite mounds are keystone structures in African savannas, affecting multiple ecosystem processes. Despite the large size of termite mounds having the potential to modify conditions around them, patterns of mound-induced ecosystem effects have been assumed to be isotropic, with little attention given to how effects might vary around mounds. We measured soil nitrogen content, grass species composition, and mammalian grazing on and off termite mounds in the four cardinal directions, and across wet and dry seasons at three savanna sites varying in mean annual rainfall in South Africa's Kruger National Park. Evidence of directional effects (anisotropy) on ecosystem properties around termite mounds varied with site. Grass species composition differed between north- and south-facing slopes at the two drier sites where mounds were taller. However, differences in grazing extent and soil nitrogen content around mounds were only present at the intermediate rainfall site where mammalian herbivore biomass was highest, and mounds were of medium height. Our results suggest that termite mound effects display significant variation with direction, but that the emergence of directional effects is context dependent. Our results further suggest that such context-dependent directional effects can lead to positive feedback loops between termites, abiotic conditions, and mammalian herbivores.