Pollination by Hemimepsis wasps: A newly described South African guild with an analysis of trait convergence between guild members

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Elephant impact on marula trees, and African honeybees as a mitigation method

A dissertation submitted to the Faculty of Science, University of the Witwatersrand, in fulfilment of the requirements for the degree of Master of Science, Johannesburg, South Africa 2017Concerns exist over the continual decline of marula trees (Sclerocarya birrea subsp. caffra) as a result of African elephant (Loxodonta africana) impact and a lack of recruitment and regeneration. One strategy of protecting adult marula trees is the usage of elephant mitigation methods. This study took place in Jejane Private Nature Reserve (JPNR), a protected area which recently opened up to the Greater Kruger National Park and had not had elephants in over 100 years. The aim of the study was to investigate the changes to the marula population structure in JPNR three years after the migration of elephants to the area due to fence removal, and to test whether African honeybees (Apis mellifera subsp. scutellata) could be used as a mitigation method for elephant impact on marula trees. A previous size-class survey had been done on a sample of JPNR’s marula population in 2009, prior to the fence removal in 2013. A resurvey of these trees was used to assess the elephant-induced impact and mortality levels on the marula trees and to compare these levels to previously recorded impact and mortality levels on marula trees in the Kruger National Park (KNP). Marula seed predation levels and seedling recruitment were also assessed to address recruitment concerns. The resurveyed marula population had declined by 23.8% post-elephant migration, with the highest annual mortality rates (AMR) and impact scores recorded for trees in the 5 - 11 m height classes. Impact scores on marula trees in JPNR were higher than impact scores recorded on KNP marula trees. Only two marula seedlings were found across all transects, with evidence of high seed predation on marula endocarps. JPNR displayed an adult-dominated marula population with a lack of regeneration, possibly due to a lack of fire which has increased available shelter for seed predators such as small mammals. African honeybees were then used to investigate their effectiveness as an elephant mitigation method and to compare this method against wire-netting (a method experimentally used to prevent ring-barking by elephants). Fifty active beehives were hung from 50 marula trees, with another 50 dummy (inactive) beehives hung from branches on the opposite ends of each beehive tree’s main stem. Fifty additional marula trees were wire-netted and a further 50 were used as control trees. Elephant impact on all 150 trees was measured prior to the addition of treatments and post-treatment addition for nine months. 54% of the control trees received some form of elephant impact, in comparison to 28% of the wire-netted trees and only 2% of the beehive trees. Wire-netting protected marula trees against bark-stripping, but did not prevent elephants from breaking branches. Beehives proved highly efficient at mitigating all forms of elephant impact. The financial cost and maintenance required for the beehive mitigation method is greater than that of wire- netting, but the beehives can provide honey and pollination services as an additional benefit. The results of this study illustrate that African honeybees can be used as an effective non-lethal mitigation method for elephant impact on marula trees and are a viable strategy to reduce human-elephant conflict in South Africa’s protected areas.MT 201

Spatio-temporal analysis of woody vegetation data at various height classes within a semi-arid savanna : insights on environmental drivers, distributions and dynamics.

Doctor of Philosophy in Biology. University of KwaZulu-Natal, Durban 2015.Savanna vegetation is regulated by bottom-up (e.g. soil and rainfall) and top-down (e.g. fire and herbivory) factors. Little is known about how these factors influence biodiversity at regional scales, particularly spatial patterns of woody plant cover, species richness, distribution and composition at various height classes. Vegetation studies are often restricted to hillslope or catchment-level scales due to time and/or logistical constraints, however there is a need to understand patterns of vegetation dynamics at regional scales as well. The main aim of this thesis was to better understand the factors (bottom-up and top-down) that influence vegetation structural dynamics in terms of species richness, distribution, composition and woody cover at regional scales. Using a rapid data collecting technique within n>1800 field sites across the entire Kruger National Park, South Africa, woody species were identified and categorized into one of three height classes; shrub (0.75m – 2.5m), brush (2.5m-5.5m) and tree (>5.5m). From this, several vegetation attributes were collated; such as species composition, richness and woody cover at each height class. A range of covariates related to disturbance (i.e. fire frequency and elephant density), landscape (e.g. aspect, altitude) and climate (temperature, rainfall) were used to identify which covariates were significantly associated with each attribute of interest. A primary finding of this research highlights the importance of bottom-up factors, such as geology and rainfall, for woody vegetation spatial distribution at regional scales. However, depending on the response variable (e.g. species richness, woody cover, species distribution), the various height classes were not influenced equally by each specific environmental factor. Overall, both climatic and non-climatic factors were highly associated with species distribution and richness, but top-down factors such as fire and herbivory were significantly associated with maintaining plant structural variation. This influenced species community composition but not species richness at each height class. Similarities in composition were related to degree of disturbance rather than the disturbance alone at specific height classes. Therefore, while species distribution and woody cover were spatially regulated by specific climatic and non-climatic factors, top-down disturbances were essential in preserving structural heterogeneity. Future vegetation monitoring campaigns in disturbance-driven systems should be cognizant of the importance of vegetation structure

Breeding and dispersal implications for the conservation of the Southern Ground Hornbill Bucorvus leadbeateri

Populations of secondary tree-cavity nesting bird species are often limited by a shortage of natural nesting sites. For the Southern Ground Hornbill Bucorvus leadbeateri that typically nests in natural tree cavities, the shortage of nesting sites is one factor potentially limiting population growth. The species is listed as endangered in South Africa, and vulnerable throughout the rest of its range. Nest boxes can improve the conservation status of threatened birds that are limited by nest-site availability. However, nest boxes or other types of artificial nests are not always beneficial to the target species, and their value as a conservation tool needs to be tested for each species. Wooden nest boxes were installed for ground hornbills in a study area in north eastern South Africa with a paucity of natural nest sites. In this thesis, I assess productivity, timing of breeding, and dispersal in the Southern Ground Hornbill in a study area supplemented with nest boxes and discuss the implications for the conservation of this endangered species. Nest boxes are an effective conservation tool to improve productivity in areas lacking natural tree cavity nesting sites. Breeding success (calculated as the proportion of nesting attempts that fledged a chick) and predation levels were similar for groups using nest boxes and natural nests. Natural nests were more buffered against cooling night temperatures, but otherwise nest boxes provided nesting conditions that were no better than natural nests. Timing of breeding for nests in natural tree cavities and nest boxes were similar. However, groups with access to a nest box attempted breeding more often than groups with access to a natural nest only, resulting in an 15 % increase in the number of fledglings per group compared to an adjacent protected area with no artificial nests. The number of breeding groups in the study area increased by 460 % over 12 years. However, there is a limit to the density of breeding groups. Breeding success was highest when breeding density was one breeding group per 90-120 km², so nests should be spaced ~10 km apart. Given that the threats to ground hornbills include persecution and poisoning, increasing the reproductive rate by providing nest boxes should assist in slowing the decline by the increased recruitment of offspring into the population. Timing of breeding varied across years. The first eggs laid each year ranged from 9 September to 14 November, and median lay date was 03 November. Breeding attempts that were initiated early in the season were more likely to fledge a chick than those initiated later in the season. Timing of breeding was delayed during warmer springs, particularly under dry conditions. In savannas, hotter spring temperatures could limit food availability, for example, if higher temperatures cause the vegetation to dry out, resulting in a rapid decline in insect flush, especially in the phytophagous insect groups that form a large part of the ground hornbill diet. Factors to consider when constructing and placing nest boxes include thickness of the cavity walls, entrance height above ground and density of nest boxes placed in the landscape. Breeding attempts in natural nests and nest boxes with thicker nest walls and those positioned with higher entrances above the ground increased breeding success. Therefore, nests should be constructed with cavity walls at least 6 cm thick and placed so that the entrances are situated > 6 m above the ground. With 186 ringed chicks fledging from the study area after the installation of nest boxes, it was possible to observe their dispersal within the study area and farther away into the adjacent Kruger National Park. There was no evidence for sex-biased dispersal. Males and females dispersed at similar ages, and over similar distances, raising interesting questions about inbreeding avoidance mechanisms in this species. If females do not disperse beyond the range of related males, how do related individuals avoid pairing, and what forms of individual recognition exist

The ecology and conservation biology of Lilian's lovebird Agapornis lilianae in Malawi.

Ph. D. University of KwaZulu-Natal, Pietermaritzburg 2014.Despite their popularity, parrots are the world’s most threatened birds. Lovebirds (Agapornis) are very popular pet and aviary birds and as ecologically specialised species in the wild, they are also among the most threatened group of birds. Lilian’s lovebird Agapornis lilianae is a mopane Colophospermum mopane woodland specialist. This study represents the first detailed investigation of the species ecology in the wild. The current distribution of Lilian’s lovebird in Malawi was explored. Furthermore, the extent of the largest resident population in Liwonde National Park (LNP) was investigated. Five new atlas records are reported; three within 40-56 km of the LNP population, and two were over 150 km south and north of LNP respectively. One of them in Kasungu National Park is about 66 km from the Lilian’s lovebird population in Luangwa Valley, Zambia. Lilian’s lovebirds occurred throughout LNP with the highest abundance in the central section. Seasonal movements to areas outside the park were recorded. A variety of vegetation types were used by the lovebirds. The strongest vegetation associations were with seasonally wet grasslands and not mopane woodlands. The abundance and density of the Lilian’s lovebird in LNP was investigated. The highest density estimates of 17 ± 4.8 lovebirds km-2 were recorded in LNP’s mopane woodland. However, number of observations per transect differed significantly. Waterhole counts had the lowest estimates (10 ± 3.5 lovebirds). Flyway counts had the intermediate estimate (14 ± 3.0 lovebirds). The total population of Lilian’s lovebirds in LNP is therefore estimated to be about 4000 individuals. The use of line transect counts at the end of the rainy season is recommended for continued monitoring of Lilian’s lovebirds abundance in LNP. Lilian’s lovebird is a secondary cavity user adapted to mopane woodlands. We investigated its roost characteristics and roosting behaviour. We quantified tree and roost site variables for roost and non-roost trees. Roosting behaviour was observed during the morning and late afternoon. Lilian’s lovebirds’ roosts were located in large tall mopane trees with a mean diameter at breast height (dbh) of 57.4 ± 1.64 m, a mean height of 16.5 ± 0.42 m, and with a mean cavity height of 10.0 ± 0.05 m. All roosts were located in mopane trees within mopane woodland with 10 – 50 % tree cover. Non-roost areas had significantly smaller trees (mean dbh = 39.4 ± 1.72 m) and were located significantly closer together. Human disturbance was low in both areas, however, evidence of elephant Loxodonta africana browsing was high with large areas of stunted mopane woodland recorded in non-roost areas. We recommend that the current LNP vegetation map be updated to highlight areas of stunted mopane woodland unsuitable for Lilian’s lovebird roosts. The impact of elephant browsing on large mopane trees should be assessed to understand its impact on the availability of suitable cavities for lovebirds and other tree cavity-reliant vertebrate species. Investigations into the diet and foraging behaviour of the Lilian’s lovebird revealed they fed on 30 different plant species. These occurred in six habitat types, two of which were outside LNP (agriculture fields and dambos). In the wet season majority of Lilian’s lovebirds (23 %) foraged in dambo areas, whilst in the dry season (August – November) the lovebirds mainly foraged in grasslands with tree cover (18 %). In mopane woodland feeding flock sizes differed significantly between the wet (mean = 20 ± 1.0 lovebirds) and dry season (mean = 34 ± 2.3 lovebirds). Grass seeds were their main food source from December to June. Lilian’s lovebirds diet was more diverse from July to November and included leaves, leaf buds, fruits, fruit seeds and herbs. Grass seeds fed on during the wet season had a high protein and energy content. The Lilian’s lovebirds foraging habitat is protected within LNP, however, early burning in areas outside the park needs to be monitored. The breeding biology of the Lilian’s lovebird was investigated. Data were collected through a combination of direct observations and infrared camera traps during three breeding seasons. Results show large similarities with the black-cheeked lovebird A. nigrigens in Zambia. The breeding season was from February to May. Lilian’s lovebirds nested mainly in south-east oriented deep cavities (≥ 1 m) located in large mopane trees (mean dbh = 57.6 ± 2.35 cm). Nests were located in loose clusters in the areas they roosted (mean distance to nearest nest = 24.2 m). Nest fidelity was observed. Clutch size ranged from 3 – 6 eggs, (mean 5.0 ± 0.22). We recorded 49 % hatching success and 69 % fledging success. Results suggest a low breeding success mainly due to the loss of eggs to predation. The use of poison to kill wildlife is a threat to biodiversity. In LNP illegal hunters poison naturally occurring waterholes to catch mammals and birds for food. Lilian’s lovebirds are among the victims at these poisoned waterholes. Lilian’s lovebird population in LNP represents about 20 % of the global population. The drinking habits of the Lilian’s lovebird, the availability of natural waterholes and the occurrence of poisoning incidents in LNP were investigated. Results showed Lilian’s lovebirds congregate at waterholes in the dry season with flock sizes ranging from 1 to 100 individuals. Significantly larger flock sizes were seen in the dry season compared with the wet season. The number of poisoning incidents/year ranged from 1 to 8. The dry season had the highest numbers of poisoning incidents. Lilian’s lovebirds were killed at approximately four poisoning incidents each year between 2000 and 2012. The number of lovebirds found dead at a poisoned pool ranged from 5 to 50 individuals. A list of other species affected by the poisoning is provided. There is need for increased efforts in preventing this lethal activity in the park. Avian diseases are considered to be one of the key threats to bird conservation. Psittacine beak and feather disease (PBFD) is the most significant infectious disease in psittacines. It is caused by the beak and feather disease virus (BFDV) and currently has no cure. PBFD threatens the survival of wild populations of endangered parrots in Africa. The occurrence of BFDV was investigated in wild populations of Lilian’s lovebird. In addition, evidence of blood parasites presence was also investigated to determine their general health. All samples (n = 48) tested negative for BFDV. Blood parasites were observed in 13 of the 48 samples (27 %). Investigation of virus occurrence in other known populations of the species is recommended to assess the conservation risk faced. Lilian’s lovebirds (n = 55) were mist-netted and ringed in LNP. Measurements showed that females were significantly larger than males. About 50 % of the birds ringed in October were half way through their primary moult indicating that moulting starts in earlier months possibly just after the breeding season in April. This study highlights three of the key threats (waterhole poisoning, habitat loss and predation) to the conservation of Lilian’s lovebirds in LNP and provides proposed actions to address these threats

African savanna elephant (Loxodonta africana) impacts on vegetation in a fenced area and the broader implications for elephant conservation.

The first chapter assessed whether a monitored population of African savanna elephants (Loxodonta africana) impacted tree species in the small, fenced Karongwe Private Game Reserve (KPGR), South Africa and to determine levels of tree recovery. Trees in high-use areas were significantly less likely to show signs of debarking and push over. Tall trees were significantly more likely to be impacted by elephants, being associated with heightened risks of debarking and branches being broken but lower risks of being pushed over. Trees close to the fence line were not more impacted than trees near the centre of the reserve. The level of use, distance to the fence and tree height were not significant predictors of tree recovery indicators. Future mitigation efforts should focus on trees with high levels of impact and low levels of recovery. The second chapter considered how elephant impact influenced secondary damages to trees and how this effected tree recovery levels. Secondary damage was identified as insects, where wood borers and termites were considered. Irrespective of tree height, termites were found to be more likely to colonise damaged trees without signs of recovery andwood borers were more likely to colonise damaged trees showing signs of recovery. Therefore, carefully considering management approaches for elephant-induced termite and wood borer impact on trees should be applied in this fenced reserve. Following this, the third chapter considered an alternative method for identifying tree height in the KPGR, using aerial photographs with 3-D photogrammetry. Initial findings here indicate that this method was successful at identifying tree height within this fenced environment. This method should be repeated in such environments in conjunction with other mapping agencies. Lastly, the fourth chapter investigated school children’s perceptions of elephants and elephant conservation at different geographical locations. Different levels of exposure to elephants were significant in influencing school children’s perceptions. More research is required to enhance environmental education practises to support elephant conservation across different locations

Seed dispersal in South African trees: with a focus on the megafaunal fruit and their dispersal agents

Includes bibliographical referencesSeed dispersal is a key process. It is important in plant population biology because it influences the fate of seeds and the probability of recruitment, in plant biogeography since dispersal mode can influence the distribution range and rate of response to environmental change and habitat fragmentation, and in animal ecology since fruits can be an important dietary item (Wang and Smith, 2002). The majority of trees in the tropics (70 – 90%) and a large proportion of trees in temperate regions (up to 60%) rely on vertebrates for their dispersal (Howe and Smallwood, 1982; Fleming et al., 1987, Willson, 1990). Vertebrate dispersers range in size from 5g mistletoe birds (Dicaeidae) to 7,500,000g elephants (Elephantidae). The range and distribution of frugivore sizes is not uniform across ecosystems or geographical regions (Mack, 1993). These differences, one might suspect would be mirrored in the range and distribution of fruit size. This is not the case; in South America where the largest frugivorous mammal is the tapir (300kg; Hansen and Galetti, 2009), there is a subset of fruit that are conspicuously large. The paradoxical existence of such large fruit in the lowlands of Costa Rica was first noted by Janzen. In collaboration with Pleistocene faunal expert Paul Martin they conjectured that these fruit were ecological anachronisms that had evolved in the presence of large terrestrial vertebrates (>1000kg - megafauna) but had remained long after their demise (Janzen and Martin, 1982)

Annals of ivory : perspectives on African elephant Loxodonta africana (Blumenbach 1797) feeding ecology from a multi-decadal record.

Includes abstract.Includes bibliographical references (leaves 202-256).This thesis explores the dietary responses of African elephants (Loxodonta africana) to environmental change by testing the hypothesis that diet switching (from predominantly browse-based to more grass-rich diets) is driven by cyclical patterns of climate and habitat change in a southern African savanna. Elephants are thought to have substantial impacts on their environments, primarily because they consume large amounts of vegetation over sustained periods. However, the woody plant composition of their diet varies considerably across space and through time, so that in some instances they have been found to be almost pure grazers. Tracking these changes by traditional approaches (e.g. field observations) is difficult because of the geographical and temporal constraints inherent to these methods. Stable light isotope tracking of diet allows diet switching to be studied over multiple space/time scales. Here, I use stable isotope data from elephant faeces, tail hair, and ivory to record short- (monthly), medium- (seasonal to annual), and long-term (decadal) ecological variability, respectively, of elephant diets in the Kruger National Park, South Africa. Results from faeces collected at monthly resolution for one year confirm findings of a previous study (based on biannually-collected samples over two years) that elephants generally consume more grass in the more wooded habitats of the northern Kruger Park, but that there is a greater degree of seasonal diet switching in southern Kruger Park habitats. Moreover, diet changes also relate to changes in underlying bedrock across Kruger Park. Isotopic time-series produced by serial profiling of tail hairs confirm patterns observed in faeces. Long-term diet histories of individuals are derived from serial isotope sampling of ivory, yielding records that represent several decades of an animal’s life, at sub-annual (seasonal) resolution. Overlaying individual ivory series in time produces the first, to my knowledge, multidecadal record of African elephant diet, dating from 1903 to 1993. Contrary to expectations, stable carbon, nitrogen, and oxygen isotope records from ivory do not correlate well with cyclical climate trends for the study region. Rather, pronounced diet shifts are observed during extreme climatic events (floods and droughts), and the greatest levels of intra- and inter-annual variability coincide with significant changes in park management policy during the 20th century, i.e. the introduction of water provision programs after the mid 1930s, and the onset of elephant population control in 1967. It is proposed that such direct intervention has played the biggest role in disturbance of elephant-plant equilibria during the 20th century, and further studies to improve our understanding of this phenomenon will be instrumental to development of appropriate management strategies for the 21st century