Conservation overstretch and long-term decline of wildlife and tourism in the Central African savannas.

When in 2010 the world's governments pledged to increase protected area coverage to 17% of the world's land surface, several Central African countries had already set aside 25% of their northern savannas for conservation. To evaluate the effectiveness of this commitment, we analyzed the results of 68 multispecies surveys conducted in the seven main savanna national parks in Central Africa (1960-2017). We also assembled information on potential drivers of large herbivore population trends (rainfall and number of rangers) and on tourist numbers and revenues. In six out of the seven parks, wild large herbivore populations declined dramatically over time, livestock numbers increased severalfold, and tourism, the pillar under a once thriving local wildlife industry, collapsed. Zakouma National Park (Chad) stood out because its large herbivore populations increased, an increase that was positively correlated with rainfall and number of rangers (a proxy for management inputs). With increasing insecurity and declining revenues, governments find themselves confronted with too few resources to protect vast areas. To deal with this conversation overstretch, we propose to extend the repeatedly promoted solutions--scaled up funding, enhanced management--with a strategic retreat, focusing scarce resources on smaller areas to save wildlife in the Central African savannas.

Sobredimensionamiento de la Conservación y la Declinación a Largo Plazo de la Fauna y el Turismo en las Sabanas de África Central Resumen Cuando los gobiernos del mundo se comprometieron en 2010 a incrementar la cobertura de áreas protegidas al 17% de la superficie terrestre del planeta, varios países del centro de África ya habían dispuesto el 25% de sus sabanas ubicadas al norte de la región para la conservación. Para evaluar la efectividad de este compromiso, analizamos los resultados de 68 censos multiespecies realizados en los siete parques nacionales principales de la sabana en África Central (1960-2017). También ensamblamos información sobre los causantes principales de las tendencias poblacionales de los grandes herbívoros (lluvias, número de guardaparques) y sobre las cifras e ingresos del turismo. En seis de los siete parques, las poblaciones de los grandes herbívoros silvestres declinaron dramáticamente con el tiempo, el número de cabezas de ganado incrementó varias veces y el turismo, el pilar de una industria faunística próspera en su momento, colapsó. El Parque Nacional Zakouma en Chad resaltó debido a que las poblaciones de herbívoros grandes incrementaron en esta localidad, un incremento que estuvo relacionado positivamente con las lluvias y el número de guardabosques (un sustituto para las aportaciones de manejo). Con el incremento en la inseguridad y la declinación de los ingresos, los gobiernos se encuentran de frente con muy pocos recursos para proteger áreas extensas. Para afrontar este sobredimensionamiento de la conservación, proponemos ampliar las soluciones que se promueven repetidamente - incrementos al financiamiento, manejo mejorado - con un repliegue estratégico, el cual enfoque los recursos escasos en las áreas más pequeñas para rescatar a la fauna de las sabanas del centro de África.

Interface processes between protected and unprotected areas: A global review and ways forward.

Land-use changes and the expansion of protected areas (PAs) have amplified the interaction between protected and unprotected areas worldwide. In this context, 'interface processes' (human-nature and cross-boundary interactions inside and around PAs) have become central to issues around the conservation of biodiversity and ecosystem services. This scientific literature review aimed to explore current knowledge and research gaps on interface processes regarding terrestrial PAs. At first, 3,515 references related to the topic were extracted through a standardized search on the Web of Science and analyzed with scientometric techniques. Next, a full-text analysis was conducted on a sample of 240 research papers. A keyword analysis revealed a wide diversity of research topics, from 'pure' ecology to sociopolitical research. We found a bias in the geographical distribution of research, with half the papers focusing on eight countries. Additionally, we found that the spatial extent of cross-boundary interactions was rarely assessed, preventing any clear delimitation of PA interactive zones. In the 240 research papers we scanned, we identified 403 processes that were studied. The ecological effects of PAs were well documented and appeared to be positive overall. In contrast, the effects of PAs on local communities were understudied and, according to the literature focusing on these, were very variable according to local contexts. Our findings highlight key research advances on interface processes, especially regarding the ecological outcomes of PAs, the influence of human activities on biodiversity, and PA governance issues. In contrast, main knowledge gaps concern the spatial extent of interactive zones, as well as the interactions between local people and conservation actions and how to promote synergies between them. While the review was limited to terrestrial PAs, its findings allow us to propose research priorities for tackling environmental and socioeconomic challenges in the face of a rapidly changing world.

Can an herbivore affect where a top predator kills its prey by modifying woody vegetation structure?

In large mammal communities, little is known about modification of interspecific interactions through habitat structure changes. We assessed the effects of African elephants (Loxodonta africana) on features of woody habitat structure that can affect predator-prey interactions. We then explored how this can influence where African lions (Panthera leo) kill their prey. Indeed, lions are stalk-and-ambush predators and habitat structure and concealment opportunities are assumed to influence their hunting success. During 2 years, in Hwange National Park, Zimbabwe, kill sites (n = 167) of GPS-collared lions were characterized (visibility distance for large mammals, distance to a potential ambush site and presence of elephant impacts). We compared characteristics of lion kill sites with characteristics of random sites (1) at a large scale (i.e. in areas intensively used by lions, n = 418) and (2) at the microhabitat scale (i.e. in the direct surrounding available habitat, < 150 m, n = 167). Elephant-impacted sites had a slightly higher visibility and a longer distance to a potential ambush site than non-impacted sites, but these relationships were characterized by a high variability. At large scale, kill sites were characterized by higher levels of elephant impacts compared to random sites. At microhabitat scale, compared to the direct nearby available habitat, kill sites were characterized by a reduced distance to a potential ambush site. We suggest a conceptual framework whereby the relative importance of habitat features and prey abundance could change upon the scale considered.

Zebra diel migrations reduce encounter risk with lions at night.

Diel migrations (DM; back and forth diel movements along an ecological gradient) undertaken by prey to avoid predators during the day have been demonstrated in many taxa in aquatic ecosystems. In terrestrial ecosystems, prey often shift between various vegetation types whose cover determine their vulnerability (i.e., likelihood of being killed when attacked). We conceptualized that in terrestrial ecosystems, DM could also occur and that the contribution of DM and shifts in vegetation cover use in reducing predation risk should depend upon the predator behaviour and the correlation between encounter risk and vulnerability across vegetation types. We further hypothesized that when the predator distribution is predictable, terrestrial prey could have evolved DM strategies taking them away from the predator when it is active or efficient. We investigated whether plains zebras Equus quagga perform DM in Hwange National Park (Zimbabwe). There, zebras can forage in large patches of open grasslands located near waterholes where they can also easily detect predators. However, they are there at high risk of encountering their main predator, lions Panthera leo, especially at night. We found out that zebras employ a DM anti-predator strategy. Zebras forage near waterholes during the day but move away from them at sunset, when lions become active. We demonstrated that this DM, occurring over a few kilometres, dramatically reduces their night-time risk of encountering lions, which generally remain close to waterholes. Zebra changes in night-time selection for vegetation cover types reduced their risk of encountering lions much less. This may arise from a trade-off between encounter risk and vulnerability across vegetation types, with zebras favouring low vulnerability once DM has reduced encounter risk. In summary, here we (a) quantify, in a terrestrial system, the effect of a predator-induced DM on the likelihood of encountering a predator and (b) distinguish the effects of the DM on encounter risk from those related to day/night changes in selection for vegetation types. We discuss how prey partition their risk between encounter risk and habitat-driven vulnerability and why it is likely critical to understand the emergence of anti-predator behavioural strategies.

Defence strategies in African savanna trees.

Southern African savannas are commonly polarised into two broad types based on plant functional types and defences; infertile savannas dominated by broad-leaved trees typically defended by nitrogen-free secondary compounds and fertile savannas dominated by fine-leaved trees defended by structural defences. In this study, we use trait and other data from 15 wooded savanna sites in Southern Africa and ask if broad-leaved and fine-leaved species dominate on nutrient-poor and nutrient-rich soils, respectively. We then test if there is there any evidence for trade-offs in chemical (i.e., condensed tannins and total polyphenols) vs. structural defences on different soil types. We did not find strong evidence for a general divide in fine- vs. broad-leaved savannas according to soil fertility, nor for a simple trade-off between chemical and structural defences. Instead, we found savanna species to cluster into three broad defence strategies: species were high in leaf N and either (A) highly defended by spines and chemicals or (B) only structurally defended, or (C) low in leaf N and chemically defended. Finally, we tested for differences in browser utilisation between soil types and among plant defence strategies and found that browsing by meso-herbivores was higher on nutrient-rich soils and targeted species from groups A and B and avoided C, while browsing by elephants was mostly not affected by soil type or defence strategy. We propose a framework that can be used as a basis for asking strategic questions that will help improve our understanding of plant defences in savannas.

Interspecific interference competition at the resource patch scale: do large herbivores spatially avoid elephants while accessing water?

Animals may anticipate and try to avoid, at some costs, physical encounters with other competitors. This may ultimately impact their foraging distribution and intake rates. Such cryptic interference competition is difficult to measure in the field, and extremely little is known at the interspecific level. We tested the hypothesis that smaller species avoid larger ones because of potential costs of interference competition and hence expected them to segregate from larger competitors at the scale of a resource patch. We assessed fine-scale spatial segregation patterns between three African herbivore species (zebra Equus quagga, kudu Tragelaphus strepsiceros and giraffe Giraffa camelopardalis) and a megaherbivore, the African elephant Loxodonta africana, at the scale of water resource patches in the semi-arid ecosystem of Hwange National Park, Zimbabwe. Nine waterholes were monitored every two weeks during the dry season of a drought year, and observational scans of the spatial distribution of all herbivores were performed every 15 min. We developed a methodological approach to analyse such fine-scale spatial data. Elephants increasingly used waterholes as the dry season progressed, as did the probability of co-occurrence and agonistic interaction with elephants for the three study species. All three species segregated from elephants at the beginning of the dry season, suggesting a spatial avoidance of elephants and the existence of costs of being close to them. However, contrarily to our expectations, herbivores did not segregate from elephants the rest of the dry season but tended to increasingly aggregate with elephants as the dry season progressed. We discuss these surprising results and the existence of a trade-off between avoidance of interspecific interference competition and other potential factors such as access to quality water, which may have relative associated costs that change with the time of the year.

Between-gender differences in vigilance do not necessarily lead to differences in foraging-vigilance tradeoffs.

When prey are time limited in their access to food, any trade-off involving time should ultimately affect their intake rate. In many herbivores, males and females experience different ecological pressures affecting their survival and reproduction because of differences in morphology, physiology and energy/nutrient requirements. If males and females have different vigilance strategies that affect their intake rates differently, they will suffer different foraging costs. This is particularly relevant in sexually monomorphic herbivores, where the two sexes have similar basal energy/nutrient requirements and risk of predation. We investigated how gender, reproductive status, age, group size, predation risk, and food biomass affected vigilance, intake rate, and their trade-off in a monomorphic species, the plains zebra (Equus quagga). Males were more vigilant than females, and lactating females were less vigilant than other females; the levels of vigilance were low (ca. 10 % of feeding time). The effects on time spent feeding, bite rates and intake rates were small and statistically not significant. Reproductive status did not affect the strength of the relationship between vigilance and intake rate, but intake rates increased with group size and, for adult females, were higher in tall grass. While gender and reproductive status were major drivers of vigilance, and group size and food biomass of the rate of food intake, males and females adjust their bite rates and food intake with vigilance in similar ways. Our results support the hypothesis that in monomorphic animals, males and females seem to make similar trade-offs (i.e. adjustments) between vigilance and intake rate.

Diet quality in a wild grazer declines under the threat of an ambush predator.

Predators influence prey populations not only through predation itself, but also indirectly through prompting changes in prey behaviour. The behavioural adjustments of prey to predation risk may carry nutritional costs, but this has seldom been studied in the wild in large mammals. Here, we studied the effects of an ambush predator, the African lion (Panthera leo), on the diet quality of plains zebras (Equus quagga) in Hwange National Park, Zimbabwe. We combined information on movements of both prey and predators, using GPS data, and measurements of faecal crude protein, an index of diet quality in the prey. Zebras which had been in close proximity to lions had a lower quality diet, showing that adjustments in behaviour when lions are within short distance carry nutritional costs. The ultimate fitness cost will depend on the frequency of predator-prey encounters and on whether bottom-up or top-down forces are more important in the prey population. Our finding is the first attempt to our knowledge to assess nutritionally mediated risk effects in a large mammalian prey species under the threat of an ambush predator, and brings support to the hypothesis that the behavioural effects of predation induce important risk effects on prey populations.

Seasonal activity patterns of a Kalahari mammal community: Trade-offs between environmental heat load and predation pressure.

Mammals in arid zones have to trade off thermal stress, predation pressure, and time spent foraging in a complex thermal landscape. We quantified the relationship between the environmental heat load and activity of a mammal community in the hot, arid Kalahari Desert. We deployed miniature black globe thermometers within the existing Snapshot Safari camera trap grid on Tswalu Kalahari Reserve, South Africa. Using the camera traps to record species' activity throughout the 24-h cycle, we quantified changes in the activity patterns of mammal species in relation to heat loads in their local environment. We compared the heat load during which species were active between two sites with differing predator guilds, one where lion (Panthera leo) biomass dominated the carnivore guild and the other where lions were absent. In the presence of lion, prey species were generally active under significantly higher heat loads, especially during the hot and dry spring. We suggest that increased foraging under high heat loads highlights the need to meet nutritional requirements while avoiding nocturnal activity when predatory pressures are high. Such a trade-off may become increasingly costly under the hotter and drier conditions predicted to become more prevalent as a result of climate change within the arid and semi-arid regions of southern Africa.

Using the multivariate Hawkes process to study interactions between multiple species from camera trap data.

Interspecific interactions can influence species' activity and movement patterns. In particular, species may avoid or attract each other through reactive responses in space and/or time. However, data and methods to study such reactive interactions have remained scarce and were generally limited to two interacting species. At this time, the deployment of camera traps opens new opportunities but adapted statistical techniques are still required to analyze interaction patterns with such data. We present the multivariate Hawkes process (MHP) and show how it can be used to analyze interactions between several species using camera trap data. Hawkes processes use flexible pairwise interaction functions, allowing us to consider asymmetries and variations over time when depicting reactive temporal interactions. After describing the theoretical foundations of the MHP, we outline how its framework can be used to study interspecific interactions with camera trap data. We design a simulation study to evaluate the performance of the MHP and of another existing method to infer interactions from camera trap-like data. We also use the MHP to infer reactive interactions from real camera trap data for five species from South African savannas (impala Aepyceros melampus, greater kudu Tragelaphus strepsiceros, lion Panthera leo, blue wildebeest Connochaetes taurinus and Burchell's zebra Equus quagga burchelli). The simulation study shows that the MHP can be used as a tool to benchmark other methods of interspecific interaction inference and that this model can reliably infer interactions when enough data are considered. The analysis of real data highlights evidence of predator avoidance by prey and herbivore-herbivore attraction. Lastly, we present the advantages and limits of the MHP and discuss how it can be improved to infer attraction/avoidance patterns more reliably. As camera traps are increasingly used, the multivariate Hawkes process provides a promising framework to decipher the complexity of interactions structuring ecological communities.

Androgen changes and flexible rutting behaviour in male giraffes.

The social organization of giraffes (Giraffa camelopardalis) imposes a high-cost reproductive strategy on bulls, which adopt a 'roving male' tactic. Our observations on wild giraffes confirm that bulls indeed have unsynchronized rut-like periods, not unlike another tropical megaherbivore, the elephant, but on a much shorter timescale. We found profound changes in male sexual and social activities at the scale of about two weeks. This so far undescribed rutting behaviour is closely correlated with changes in androgen concentrations and appears to be driven by them. The short time scale of the changes in sexual and social activity may explain why dominance and reproductive status in male giraffe in the field seem to be unstable.

Insights for integrated conservation from attitudes of people toward protected areas near Hwange National Park, Zimbabwe.

Increase in human settlements at the edge of protected areas (PAs) is perceived as a major threat to conservation of biodiversity. Although it is crucial to integrate the interests of surrounding communities into PA management, key drivers of changes in local populations and the effects of conservation on local livelihoods and perceptions remain poorly understood. We assessed population changes from 1990 to 2010 in 9 villages located between 2 PAs with different management policies (access to natural resources or not). We conducted semi-directive interviews at the household level (n =217) to document reasons for settlement in the area and villager's attitudes toward the PAs. We examined drivers of these attitudes relative to household typology, feelings about conservation, and concerns for the future with mixed linear models. Population increased by 61% from 2000 to 2010, a period of political and economic crisis in Zimbabwe. Forty-seven percent of immigrants were attracted by the area; others had been resettled from other villages or were returning to family lands. Attitudes toward PAs were generally positive, but immigrants attracted by the area and who used resources within the PA with fewer restrictions expressed more negative attitudes toward PAs. Household location, losses due to wild animals, and restrictions on access to natural resources were the main drivers of this negative attitude. Profit-seeking migrants did not expect these constraints and were particularly concerned with local overpopulation and access to natural resources. To avoid socio-ecological traps near PAs (i.e., unforeseen reduced adaptive capacity) integrated conservation should address mismatches between management policy and local expectations. This requires accounting for endogenous processes, for example, local socio-ecological dynamics and values that shape the coexistence between humans and wildlife.

Validation of a field-friendly faeces drying and storage method for quantifying faecal glucocorticoid metabolites in African elephants (Loxodonta africana) opens up new perspectives for conservationists.

Faecal glucocorticoid metabolites (fGCMs) are a relevant means of non-invasively assessing adrenocortical activity and thus, a key physiological stress response in wildlife populations. However, the widespread use of fGCMs as a stress-related biomarker in conservation biology is often hampered by the logistical challenge of storing collected faecal material frozen until it reaches the laboratory for analysis. Although alternative approaches to minimize potential alteration of fGCM composition post-defecation have been recently identified, there is to our knowledge, no satisfactory alternative method established for the preservation of elephant dung. In this study, we validated a field-friendly protocol for dehydrating African elephant faeces samples using a food dehydrator with desiccant and investigated the stability of fGCM concentrations in the dehydrated faeces when stored at ambient temperature. We collected 40 faecal samples from African elephants and compared fGCM concentrations of freeze-dried and dehydrated sample sub-sets. Samples dried in the field showed a slight but significant overall -6% reduction in fGCM concentration compared with frozen control samples. However, fGCM concentrations following field dehydration protocol match those of control samples with high accuracy, as evidenced by the low bias and strong coefficient of determination between the two approaches (R2 = 0.88). In addition, over nearly 2 months, storage time at ambient temperature of the dehydrated samples had no effect on the fGCM concentrations compared with those measured in the control samples (F-statistic = 1.82; P = 0.18). Dehydrating the samples in the field thus provides an easy and cost-effective alternative to freezing, especially when working in remote areas with unstable electrical supply. Our results encourage the widespread use of fGCMs by conservationists as non-invasive means of steroid monitoring of African elephants in the current context of a general increase in wildlife welfare research. Future studies are needed to extend the use of this protocol to other species and to other steroid classes.

Less bins, less baboons: reducing access to anthropogenic food effectively decreases the urban foraging behavior of a troop of chacma baboons (Papio hamadryas ursinus) in a peri-urban area.

In South Africa, chacma baboons (Papio hamadryas ursinus) living near peri-urban areas may forage on anthropogenic food. Baboons have been recorded to damage crops, scatter waste from trash bins, and damage homes. A number of methods have been tested over the past 20 years to solve these problems, but none proved successful over the long-term or involved considerable costs. An efficient management system requires a detailed knowledge of how baboon troops proceed and organize during these urban foraging actions. This study examines the response of a troop of baboons to an experimental reduction of anthropogenic food sources in a peri-urban environment, the George campus of Nelson Mandela University (SA). We gradually suppressed access to waste food in trash cans, reducing the amount of anthropogenic food available. This change in food availability led baboons to modify their urban foraging strategy. They compensated for the lack of anthropogenic food by spending more time foraging on natural food and less time in urban areas. However, the troop still exploited waste-free areas during the experiment and even more when the conditions were normal again. Overall, these results show the ability of baboons to adapt to changes in anthropogenic food availability but also that they are highly dependent on this type of resource. Limiting its access is a mitigation strategy that humans must absolutely develop for reaching a high level of coexistence with baboons.

Considerations on effort, precision and accuracy for long-term monitoring of African lions (Panthera leo), when using Bayesian spatial explicit capture-recapture models, in fenced protected areas.

Intensive management is frequently required in fenced wildlife areas to reduce deleterious effects of isolation. Decisions on how best to manage such wildlife are ideally informed by regular and reliable estimates of spatiotemporal fluctuations in population size and structure. However, even in small, fenced areas, it is difficult and costly to regularly monitor key species using advanced methods. This is particularly the case for large carnivores, which typically occur at low density and are elusive yet are central to management decision-making due to their top-down effects in ecosystems and attracting tourism. In this study, we aimed to provide robust estimates of population parameters for African lions (Panthera leo) and use the data to inform a resource-efficient long-term monitoring programme. To achieve this, we used unstructured spatial sampling to collect data on lions in Pilanesberg National Park, a small (~550 km2) fenced protected area in South Africa. We used Bayesian spatial capture-recapture models to estimate density, abundance, sex ratio and home range size of lions over the age of 1 year. Finally, to provide guidance on resource requirements for regular monitoring, we rarefied our empirical data set incrementally and analysed the subsets. Lion density was estimated to be 8.8 per 100 km2 (posterior SD = 0.6), which was lower than anticipated by park management. Sex ratio was estimated close to parity (0.9♀:1♂), consistent with emerging evidence in fenced lion populations, yet discordant with unfenced populations, which are usually ~2♀:1♂ in healthy, source populations. Our rarefied data suggest that a minimum of 4000 km search effort needs to be invested in future monitoring to obtain accurate and precise estimates, while assuming similar detection rates. This study demonstrates an important utility of Bayesian spatial explicit capture-recapture methods for obtaining robust estimates of lion densities and other important parameters in fence-protected areas to inform decision-making.

Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna.

Vigilance allows individuals to escape from predators, but it also reduces time for other activities which determine fitness, in particular resource acquisition. The principles determining how prey trade time between the detection of predators and food acquisition are not fully understood, particularly in herbivores because of many potential confounding factors (such as group size), and the ability of these animals to be vigilant while handling food. We designed a fertilization experiment to manipulate the quality of resources, and compared awareness (distinguishing apprehensive foraging and vigilance) of wild impalas (Aepyceros melampus) foraging on patches of different grass height and quality in a wilderness area with a full community of predators. While handling food, these animals can allocate time to other functions. The impalas were aware of their environment less often when on good food patches and when the grass was short. The animals spent more time in apprehensive foraging when grass was tall, and no other variable affected apprehensive behavior. The probability of exhibiting a vigilance posture decreased with group size. The interaction between grass height and patch enrichment also affected the time spent in vigilance, suggesting that resource quality was the main driver when visibility is good, and the risk of predation the main driver when the risk is high. We discuss various possible mechanisms underlying the perception of predation risk: foraging strategy, opportunities for scrounging, and inter-individual interference. Overall, this experiment shows that improving patch quality modifies the trade-off between vigilance and foraging in favor of feeding, but vigilance remains ultimately driven by the visibility of predators by foragers within their feeding patches.

Factors that influence Cape fur seal predation on Cape gannets at Lambert's Bay, South Africa.

Seabird populations experience predation that can impact their breeding density and breeding success. The Cape gannet Morus capensis is endemic to the Benguela upwelling ecosystem and is classified as Endangered by the IUCN. They are affected by several threats, including predation by the Cape fur seal Arctocephalus pusillus pusillus. Many fledglings succumb to predation during their maiden flight across waters around the island. To curb predation, the selective culling of individual predatory seals was implemented in 2014, 2015, and 2018. Our first study objective was to determine if selective culling of Cape fur seals significantly reduced predation probability on Cape gannets. We tested whether predation probability in 2014, 2015, and 2018 was affected by fish biomass, gannet fledgling numbers, and/or the presence/absence of selective culling. Our second objective was to determine what led to fluctuations in Cape fur seal predation on Cape gannet fledglings between 2007 and 2018. We tested whether fish biomass and the amount of Cape gannet fledglings in the water affected predation probability on the fledglings. Results indicated that selective culling reduced predation within years. We found that with both increased fledgling numbers and increased fish biomass, seal predation probability was reduced. This suggests that a sustainable way to promote the conservation of Cape gannets would be to increase food availability for both the Cape fur seals and Cape gannets. Our findings, collectively with the global trend of the declining Cape gannet population and their endemism, provide reasons advocating for the conservation of the food resources of both the Cape fur seal and the Cape gannet in the Benguela system.

Simultaneous modeling of habitat suitability, occupancy, and relative abundance: African elephants in Zimbabwe.

The recent development of statistical models such as dynamic site occupancy models provides the opportunity to address fairly complex management and conservation problems with relatively simple models. However, surprisingly few empirical studies have simultaneously modeled habitat suitability and occupancy status of organisms over large landscapes for management purposes. Joint modeling of these components is particularly important in the context of management of wild populations, as it provides a more coherent framework to investigate the population dynamics of organisms in space and time for the application of management decision tools. We applied such an approach to the study of water hole use by African elephants in Hwange National Park, Zimbabwe. Here we show how such methodology may be implemented and derive estimates of annual transition probabilities among three dry-season states for water holes: (1) unsuitable state (dry water holes with no elephants); (2) suitable state (water hole with water) with low abundance of elephants; and (3) suitable state with high abundance of elephants. We found that annual rainfall and the number of neighboring water holes influenced the transition probabilities among these three states. Because of an increase in elephant densities in the park during the study period, we also found that transition probabilities from low abundance to high abundance states increased over time. The application of the joint habitat-occupancy models provides a coherent framework to examine how habitat suitability and factors that affect habitat suitability influence the distribution and abundance of organisms. We discuss how these simple models can further be used to apply structured decision-making tools in order to derive decisions that are optimal relative to specified management objectives. The modeling framework presented in this paper should be applicable to a wide range of existing data sets and should help to address important ecological, conservation, and management problems that deal with occupancy, relative abundance, and habitat suitability.

Fire weather effects on flammability of indigenous and invasive alien plants in coastal fynbos and thicket shrublands (Cape Floristic Region).

BACKGROUND: Globally, and in the Cape Floristic Region of South Africa, extreme fires have become more common in recent years. Such fires pose societal and ecological threats and have inter alia been attributed to climate change and modification of fuels due to alien plant invasions. Understanding the flammability of different types of indigenous and invasive alien vegetation is essential to develop fire risk prevention and mitigation strategies. We assessed the flammability of 30 species of indigenous and invasive alien plants commonly occurring in coastal fynbos and thicket shrublands in relation to varying fire weather conditions. METHODS: Fresh plant shoots were sampled and burnt experimentally across diverse fire weather conditions to measure flammability in relation to fire weather conditions, live fuel moisture, fuel load and vegetation grouping (fynbos, thicket and invasive alien plants). Flammability measures considered were: burn intensity, completeness of burn, time-to-ignition, and the likelihood of spontaneous ignition. We also investigated whether the drying of plant shoots (simulating drought conditions) differentially affected the flammability of vegetation groups. RESULTS: Fire weather conditions enhanced all measures of flammability, whereas live fuel moisture reduced burn intensity and completeness of burn. Live fuel moisture was not significantly correlated with fire weather, suggesting that the mechanism through which fire weather enhances flammability is not live fuel moisture. It furthermore implies that the importance of live fuel moisture for flammability of evergreen shrublands rests on inter-specific and inter-vegetation type differences in fuel moisture, rather than short-term intra-specific fluctuation in live fuel moisture in response to weather conditions. Fuel load significantly increased burn intensity, while reducing ignitability. Although fire weather, live fuel moisture, and fuel load had significant effects on flammability measures, vegetation and species differences accounted for most of the variation. Flammability was generally highest in invasive alien plants, intermediate in fynbos, and lowest in thicket. Fynbos ignited rapidly and burnt completely, whereas thicket was slow to ignite and burnt incompletely. Invasive alien plants were slow to ignite, but burnt with the highest intensity, potentially due to volatile organic composition. The drying of samples resulted in increases in all measures of flammability that were comparable among vegetation groups. Flammability, and by implication fire risk, should thus not increase disproportionately in one vegetation group compared to another under drought conditions-unless the production of dead fuels is disproportionate among vegetation groups. Thus, we suggest that the dead:live fuel ratio is a potentially useful indicator of flammability of evergreen shrublands and that proxies for this ratio need to be investigated for incorporation into fire danger indices.

Are fission-fusion dynamics consistent among populations? A large-scale study with Cape buffalo.

Fission-fusion dynamics allow animals to manage costs and benefits of group living by adjusting group size. The degree of intraspecific variation in fission-fusion dynamics across the geographical range is poorly known. During 2008-2016, 38 adult female Cape buffalo were equipped with GPS collars in three populations located in different protected areas (Gonarezhou National Park and Hwange National Park, Zimbabwe; Kruger National Park, South Africa) to investigate the patterns and environmental drivers of fission-fusion dynamics among populations. We estimated home range overlap and fission and fusion events between Cape buffalo dyads. We investigated the temporal dynamics of both events at daily and seasonal scales and examined the influence of habitat and distance to water on event location. Fission-fusion dynamics were generally consistent across populations: Fission and fusion periods lasted on average between less than one day and three days. However, we found seasonal differences in the underlying patterns of fission and fusion, which point out the likely influence of resource availability and distribution in time on group dynamics: During the wet season, Cape buffalo split and associated more frequently and were in the same or in a different subgroup for shorter periods. Cape buffalo subgroups were more likely to merge than to split in open areas located near water, but overall vegetation and distance to water were very poor predictors of where fission and fusion events occurred. This study is one of the first to quantify fission-fusion dynamics in a single species across several populations with a common methodology, thus robustly questioning the behavioral flexibility of fission-fusion dynamics among environments.