The Aardvark as an ecological engineer in the Eastern Karoo: dig patterns and emergent processes

Species which play particularly important roles in ecosystem patterns and processes are recognised as keystone species or ecosystem engineers. The conservation of these species is critical for the maintenance of the ecological role they play in ecosystem function. The aardvark, Orycteropus afer, is known to play a role in biopedturbation through digging for refuges and for prey, and as a consequence of these digging activities is expected to serve the role of an ecosystem engineer and keystone species. This study aimed to describe and quantify the ecological patterns and processes driven by the aardvark through their digging activities, within the eastern Karoo, and thereby evaluate the significance of aardvarks as ecosystem engineers and keystone species in a semi-arid environment. The landscape pattern (dig abundance and distribution) and soil displacement of aardvark forage and refuge digs in a semi-arid Karoo landscape, as well as the ecological processes (generating fertile, seed retention hotspots) driven by aardvark forage digs was measured. Refuge digs occurred in the highest densities in areas characterised by a specific suite of soil, vegetation and elevation features, while forage digs occurred in high abundance in most habitats measured, and were likely driven by prey availability. Forage digs served as litter and seed retention hotspots, with accelerated decomposition rates of litter as a function of increased soil moisture in digs. The functional processes (i.e. resource capture, shelter for plants and animals, germination sites, soil aeration, organic turnover, mineralization rates, fertile soil displacement and transport) driven by aardvark digs are not restricted to refuge digs. Forage digs occurred in most habitats measured and their functional role is unique and perhaps greater across the landscape in comparison to refuge digs. The aardvark has a disproportionate effect in ecosystems, in terms of its refuge and forage digs relative to its abundance and therefore its role as a keystone species and ecosystem engineer is confirmed. Conservation and protection of this species is therefore essential to maintain its important role in ecosystem function

Megaherbivores in succulent thicket: resource use and implications

This study aims to develop a predictive understanding of the resource use, impacts and interactions of elephant Loxodonta africana and black rhinoceros Diceros bicornis in the succulent thickets of the Eastern Cape, South Africa. While these megaherbivores typically dominate the biomass, elephant are more abundant, such that their impacts off-set that of all other herbivores. Consequently, this thesis has three main foci: first, developing a mechanistic understanding of the influences of elephant; second, developing predictive insights into elephant impacts on plant communities; finally, an understanding of the knockon-effects of the impacts for coexisting rhinoceros. Thus, by documenting the diet and dietary preferences of elephant, I firstly show that only about 18 percent of the species previously thought vulnerable to herbivory, occur in the diet. This refutes the generally held belief that elephant herbivory is the primary driver of decline among plants, and emphasizes the likely contribution of other mechanisms (e.g. trampling, knock-on-effects, etc.). Thus, the accurate prediction of the impacts caused by elephant requires an understanding of previously marginalized mechanisms. From here, I quantify >50 years of impacts on the thicket shrub community and test their spatial and temporal extent near water. I confirm the vulnerability of thicket to transformation (particularly near water) as the accumulated influences of elephant reduce community composition and structure, and predict that these impacts will eventually bring about landscape-level degradation and a significant loss of biodiversity. Importantly, results show an uneven distribution of effects between elements of this community: from community composition and structure, to the structure of individual canopy species and ecological functioning. While these findings confound our interpretation of the extent of the impacts, it demonstrates the importance of explicitly recognizing biodiversity and heterogeneity for the conservation management of elephant. Finally, I test the consequences of the impacts for coexisting rhinoceros. While I show that this causes rhinoceros to change their foraging strategies in the presence of elephant at high densities, I also show that elephant may facilitate access to food for rhinoceros at reduced densities. These findings indicate the importance of elephant in driving the structure and composition of the thicket shrub community and the consequences of this for coexisting large herbivores. Thus, developing a predictive understanding of the spatial and temporal variations of elephant impacts between elements of biodiversity and the mechanisms driving these changes are key to their management. This implies that the effective conservation management of elephant can only be achieved through the careful, scientific design of monitoring programmes

Megaherbivores in succulent thicket: resource use and implications

This study aims to develop a predictive understanding of the resource use, impacts and interactions of elephant Loxodonta africana and black rhinoceros Diceros bicornis in the succulent thickets of the Eastern Cape, South Africa. While these megaherbivores typically dominate the biomass, elephant are more abundant, such that their impacts off-set that of all other herbivores. Consequently, this thesis has three main foci: first, developing a mechanistic understanding of the influences of elephant; second, developing predictive insights into elephant impacts on plant communities; finally, an understanding of the knockon-effects of the impacts for coexisting rhinoceros. Thus, by documenting the diet and dietary preferences of elephant, I firstly show that only about 18 percent of the species previously thought vulnerable to herbivory, occur in the diet. This refutes the generally held belief that elephant herbivory is the primary driver of decline among plants, and emphasizes the likely contribution of other mechanisms (e.g. trampling, knock-on-effects, etc.). Thus, the accurate prediction of the impacts caused by elephant requires an understanding of previously marginalized mechanisms. From here, I quantify >50 years of impacts on the thicket shrub community and test their spatial and temporal extent near water. I confirm the vulnerability of thicket to transformation (particularly near water) as the accumulated influences of elephant reduce community composition and structure, and predict that these impacts will eventually bring about landscape-level degradation and a significant loss of biodiversity. Importantly, results show an uneven distribution of effects between elements of this community: from community composition and structure, to the structure of individual canopy species and ecological functioning. While these findings confound our interpretation of the extent of the impacts, it demonstrates the importance of explicitly recognizing biodiversity and heterogeneity for the conservation management of elephant. Finally, I test the consequences of the impacts for coexisting rhinoceros. While I show that this causes rhinoceros to change their foraging strategies in the presence of elephant at high densities, I also show that elephant may facilitate access to food for rhinoceros at reduced densities. These findings indicate the importance of elephant in driving the structure and composition of the thicket shrub community and the consequences of this for coexisting large herbivores. Thus, developing a predictive understanding of the spatial and temporal variations of elephant impacts between elements of biodiversity and the mechanisms driving these changes are key to their management. This implies that the effective conservation management of elephant can only be achieved through the careful, scientific design of monitoring programmes

Macro-nutritional balancing in a circumpolar boreal ruminant under winter conditions

1. Differences in botanical diet compositions among a large number of moose faecal samples collected during winter correlated with the nutritional differences identified in the same samples (Mantel r = 0.89, p = 0.001), but the nutritional differences were significantly smaller (p 0).3. Available protein (AP) and total non-structural carbohydrates (TNC) were significantly correlated in observed diets but not in hypothetical diets based on food availability.4. The level of Acetoacetate in moose serum (i.e. "starvation') was weakly negatively associated with digestibility of diets (p = 0.08) and unrelated to increasing AP:TNC and AP:NDF ratios in diets (p > 0.1).5. Our study is the first to demonstrate complementary feeding in free-ranging moose to attain a nutritional target that has previously been suggested in a feeding trial with captive moose. Our results add support to the hypothesis of nutritional balancing as a driver in the nutritional strategy of moose with implications for both the management of moose and food resources

Macro-nutritional balancing in a circumpolar boreal ruminant under winter conditions

Differences in botanical diet compositions among a large number of moose faecal samples collected during winter correlated with the nutritional differences identified in the same samples (Mantel-r = 0.89, p = 0.001), but the nutritional differences were significantly smaller (p  Nutritional geometry revealed that moose mixed Scots pine Pinus sylvestris and Vaccinium spp. as nutritionally complementary foods to reach a nutritional target resembling Salix spp. twigs, and selected for Salix spp. browse (Jacob's D > 0). Available protein (AP) and total non-structural carbohydrates (TNC) were significantly correlated in observed diets but not in hypothetical diets based on food availability. The level of Acetoacetate in moose serum (i.e. ‘starvation’) was weakly negatively associated with digestibility of diets (p = 0.08) and unrelated to increasing AP:TNC and AP:NDF ratios in diets (p > 0.1). Our study is the first to demonstrate complementary feeding in free-ranging moose to attain a nutritional target that has previously been suggested in a feeding trial with captive moose. Our results add support to the hypothesis of nutritional balancing as a driver in the nutritional strategy of moose with implications for both the management of moose and food resources. </ol

Megaherbivores in succulent thicket : resource use and implications

This study aims to develop a predictive understanding of the resource use, impacts and interactions of elephant Loxodonta africana and black rhinoceros Diceros bicornis in the succulent thickets of the Eastern Cape, South Africa. While these megaherbivores typically dominate the biomass, elephant are more abundant, such that their impacts off-set that of all other herbivores. Consequently, this thesis has three main foci: first, developing a mechanistic understanding of the influences of elephant; second, developing predictive insights into elephant impacts on plant communities; finally, an understanding of the knockon-effects of the impacts for coexisting rhinoceros. Thus, by documenting the diet and dietary preferences of elephant, I firstly show that only about 18 percent of the species previously thought vulnerable to herbivory, occur in the diet. This refutes the generally held belief that elephant herbivory is the primary driver of decline among plants, and emphasizes the likely contribution of other mechanisms (e.g. trampling, knock-on-effects, etc.). Thus, the accurate prediction of the impacts caused by elephant requires an understanding of previously marginalized mechanisms. From here, I quantify >50 years of impacts on the thicket shrub community and test their spatial and temporal extent near water. I confirm the vulnerability of thicket to transformation (particularly near water) as the accumulated influences of elephant reduce community composition and structure, and predict that these impacts will eventually bring about landscape-level degradation and a significant loss of biodiversity. Importantly, results show an uneven distribution of effects between elements of this community: from community composition and structure, to the structure of individual canopy species and ecological functioning. While these findings confound our interpretation of the extent of the impacts, it demonstrates the importance of explicitly recognizing biodiversity and heterogeneity for the conservation management of elephant. Finally, I test the consequences of the impacts for coexisting rhinoceros. While I show that this causes rhinoceros to change their foraging strategies in the presence of elephant at high densities, I also show that elephant may facilitate access to food for rhinoceros at reduced densities. These findings indicate the importance of elephant in driving the structure and composition of the thicket shrub community and the consequences of this for coexisting large herbivores. Thus, developing a predictive understanding of the spatial and temporal variations of elephant impacts between elements of biodiversity and the mechanisms driving these changes are key to their management. This implies that the effective conservation management of elephant can only be achieved through the careful, scientific design of monitoring programmes

Settling the browser–grazer debate for African buffalo in grass-limited Eastern Cape thicket, South Africa

Despite extensive evidence that African buffalo Syncerus caffer are grazers, De Graaff et al. using rumen content analysis of animals that had starved to death proposed that buffalo in grass-limited Eastern Cape thicket should be considered browsers. Although these anomalous findings were initially accepted, but later challenged, the browse-dominated diet continues to be used as a foundation for hypotheses on the diet of healthy animals. Consequently, the debate around buffalo as browsers or grazers in thicket has not yet been settled. We describe the diet of buffalo in the Addo Elephant National Park and include data from other published work from the region to test the importance of grass in buffalo diet. We show that the diet is dominated by grasses, even in grass-limited thicket, and that browse species are seldom dominant foods. Thus, there is no empirical evidence to corroborate the notion that buffalo switch their diet to browse when grass availability is low. In an attempt to advance our understanding of buffalo foraging in thicket, we reiterate that De Graaff’s work is not a valid measure of buffalo diet in succulent thicket and that additional testing of the browser–grazer hypothesis is not needed. Conservation implications: Our results confirm that buffalo are grazers, rather than browsers, in grass-limited Eastern Cape thicket. Thus, additional testing of the browser–grazer hypothesis for buffalo in the region is not needed

Impact of elephant on two woody trees, <i>Boscia oleoides</i> and <i>Pappea capensis</i>, in an arid thicket-Nama Karoo mosaic, Greater Addo Elephant National Park

Despite extensive evidence of the influences of elephant on woody trees in savannah habitats, effects on trees in the succulent thickets of the Eastern Cape are relatively poorly described. Our study investigates the role and intensity of elephant impacts on Pappea capensis and the relatively rare Boscia oleoides in an arid thicket-Nama Karoo mosaic habitat of the Greater Addo Elephant National Park. We show that roughly 19% of the B. oleoides and nearly half of the P. capensis individuals recorded showed signs of elephant impact. Elephant often toppled our study trees, and where these individuals were uprooted, mortalities occurred: B. oleoides ~ 44% of the impacted trees (4 individuals); P. capensis ~ 22% of the impacted trees (29 individuals). Conservation implications: Whilst this study is restricted by limited spatial and temporal replication, P. capensis mortalities caused by elephant occurred at a rate exceeding that of other processes. Our results provide insight into the severity of the measured changes and the need to reduce the impacts. However, it would be critically important to establish the specific driver of elephant–tree interactions before any management intervention is implemented