No description available.

Our understanding of bird pollination systems has changed dramatically in the last few years. A long-standing paradigm was that hummingbirds and passerine birds select for different nectar properties in flowers (phylogenetic hypothesis). However, specialist passerines, such as sunbirds, have similar nectar preferences to hummingbirds and nectar in plants pollinated by these two bird groups is strongly convergent. Thus, as an alternative to the existing paradigm, it has been argued that the most useful distinction that can be drawn is that between specialist and generalist avian nectarivores (feeding niche hypothesis). This was supported by phylogenetically-controlled analyses that show that nectar in plants pollinated by specialist avian nectarivores (whether hummingbird or passerine) tends to have a lower volume, higher concentration and higher sucrose content than that in plants pollinated by generalist avian nectarivores. The aim of this thesis was to determine if these trends can be explained by the nectar preferences of avian nectarivores, and to determine whether the generalist-specialist dichotomy can be applied to Kniphofia, a largely bird-pollinated African plant genus. This thesis consists of two sections. In the first, I present data from equicaloric choice experiments to determine sugar preferences of both specialist and occasional nectar feeding birds across a range of concentrations. In addition, I determine apparent sugar assimilation efficiencies and concentration preferences for a range of occasional nectar feeding birds. Using Malachite Sunbirds (Nectarinia famosa), Dark-capped bulbuls (Pycnonotus tricolor), Speckled Mousebirds (Colius striatus) and Red-winged Starlings (Onychognathus morio) (plus Village Weavers (Ploceus cucullatus) in an appendix) as representatives of the major groups of nectar feeding birds in South Africa, I show that some differences occur between occasional nectar feeding birds and all specialist nectar feeding birds studied so far. Specialists show a switch from hexose preference at low (5%) concentrations, exhibit no preference at medium concentrations (10-20%) and prefer sucrose or show no preference at high concentrations (25%). However, specialists are unable to maintain energy balance at low concentrations, and always select the higher concentration when given choices within this range. Occasional nectar feeding birds prefer hexose solutions at low and medium concentrations (5 and 10%), but show varied preferences at higher concentrations. Occasional nectar feeding birds are able to maintain energy balance at low concentrations, and either show no concentration preference, or a preference for lower concentrations when given a choice. Occasional nectar feeding birds show a range in ability to digest sucrose, although some species, like the Dark-capped Bulbul, are quite proficient at it. Surprisingly, no significant difference is found between specialist and occasional nectarivores sugar preferences when analysed globally, even when phyllogeny is accounted for. Instead, I found a significant relationship between body size and bird sugar preference at different concentrations, from which I build the body size hypothesis, which I suggest is a better predictor to use than bird diet type (specialist or occasional nectarivores – feeding niche hypothesis). In the second section of the thesis, I examine the associations between plant traits and nectarivore nectar preferences. This section focuses on flower morphology and nectar characteristics in Kniphofia species, and ecotypes within species, pollinated by specialist versus occasional avian nectarivores. I show that apart from sugar type, which appears to be phylogenetically constrained, flower morphology and nectar characteristics appear to diverge according to whether plants are pollinated by generalist or specialist nectar feeding birds. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2009.

Animal community structures, life histories and individual foraging behaviour are all an outcome of a trade-off between competition for resources (and thus the risk of starvation) and survival (and thus the risk of predation). The relative importance of these factors however, differs between ecosystems, and especially when comparing temperate to tropical ones, we usually find marked differences. The seasonality of tropical ecosystems is much reduced compared to temperate ones, and weather conditions are less extreme. Accordingly tropical systems are characterised by higher species diversity, and different life history traits have been found between temperate and tropical birds. However, how the different environmental factors interact, and how predation and starvation risk vary to cause these differences still remains largely unknown. We studied the feeding behaviour of several granivorous Estrildid finches in scrub savannah habitat in central Nigeria to test how they respond to varying degrees of starvation and predation risk. During field observations and aviary experiments we investigated whether there is seasonal variation in the birds’ foraging behaviour correlating with the abundance of grass seeds and tested how they respond to different group sizes and differing distances from cover (representing a difference in predation risk). Further we also carried out field observation on the natural feeding behaviour of several closely related sympartic Estrildid finches to investigate inter-specific and seasonal differences in competition and microhabitat choice to see if this could explain their coexistence. Finally we studied habitat choice, movement behaviour and breeding biology of the potentially threatened endemic Rock Firefinch (Lagonosticta sanguinodorsalis) between the wet and the dry season via radio-tracking to establish its habitat requirements and gain the first information in its life history traits and population trends. We found little seasonal variation in the species’ foraging behaviour, and parameters that varied did not do so in a consistent manner. Thus, we found little evidence for a seasonal change in the risk of starvation. However, the abundance of several bird species varied widely between seasons and species leaving during periods of food shortage might have released competition for remaining resources. Birds did not show a strong response in their feeding behaviour with respect to cover in either intake rate or timing of feeding. However, intake rate increased with group size, which we believe to be due to scramble competition rather than risk dilution. We therefore conclude that predation did not shape the foraging behaviour of tropical granivorous passerines as markedly as that of temperate ones. Rock Firefinches were found to breed between the late rainy and the early dry season. They selected inselberg habitat, where most nests were found between rocky boulders. During the dry season, when water sources in inselberg habitat had dried out, they had to fly distances of up to 700 m to the gallery forest to get water and this led to the inclusion of more scrub savannah and gallery forest within their home ranges. Daily egg survival was 0.89 ± 0.03 calculated after the Mayfield analysis and most failing nests were depredated probably mainly by lizards. We suggest that in addition to nest predation, water availability might limit breeding time and thus reproductive output of Rock Firefinches. Predation risk did not seem to be of high importance in shaping the birds’ feeding behaviour because there was no seasonal variation in the risk of starvation. We found some suggestive evidence that competition might be important and it is likely that bird populations constantly stay close to carrying capacity. In contrast to temperate regions the need to conserve water might be of higher importance in shaping the birds’ feeding behaviour. High adult survival rates might be due to reduced seasonality in the risk of starvation thereby leading to reduced predation risk on adult birds. High nest predation might also be of higher importance in shaping the life history traits of tropical passerines, but at present this suggestion remains speculative. The results fit into the general framework that there is a trade-off between starvation and predation risk, and in the absence of starvation risk for some species in tropical areas, predation risk is also relatively unimportant.

It has been suggested that fruit features such as nutrient content, size and colour have co-evolved with dispersal agent behaviour, physiology and morphology. Avian nectarivore feeding ecology is relatively well studied; however, less is known about fruit selection in avian frugivores. Previous work highlights the importance of individual factors that contribute towards fruit preferences, but few studies bring these factors together. Consequently the aim of this dissertation was to attempt this in terms of frugivory by investigating behavioural, physiological and morphological aspects of fruit selection in generalist avian dispersers. This was achieved by manipulating the nutritional content, size and colour of fruits (artificial fruits) under controlled conditions. The first part of the dissertation addresses physiological aspects of fruit selection in Red-winged Starling Onychognathus morio, Speckled Mousebird Colius striatus and Dark-Capped Bulbul Pycnonotus tricolor. Assimilation efficiency of birds fed glucose and sucrose diet treatments of varying concentration was observed. All study species showed high apparent assimilation efficiency irrespective of artificial fruit sugar concentration and type (with the exception of Red-winged Starlings on an all sucrose diet). The second part of the dissertation addresses behavioural aspects of fruit selection in the same three species by observing selective preferences between glucose and sucrose fruits of varying concentration and molarity. While Mousebirds displayed no preference for any of the diet treatments, Bulbuls occasionally favoured glucose diets over sucrose diets and Starlings always favoured glucose diets over sucrose diets. Another behavioural aspect of fruit selection was addressed in the third part of the dissertation. Colour preferences of Red-winged Starling and Speckled Mousebird were observed. Although study species did not conform to trends by favouring black and red fruits, they did show avoidance of green fruits and (Starlings) presented evidence of learning. A morphological aspect of fruit selection is presented in the fourth part of the dissertation. Starlings, Bulbuls and Mousebird beak morphology was measured to investigate if this has an effect on fruit size preferences. Starlings with the largest beak dimensions were more selective of fruit size classes than Bulbuls and Mousebirds which displayed the importance of feeding method (thrashing/swallowing/mashing) to compensate for larger fruit sizes. The final section of this dissertation is a synthesis of the observed behavioural, physiological and morphological aspects of fruit selection in Red-winged Starlings, Dark-capped Bulbuls and Speckled-Mousebirds. / Thesis (M.Sc.)-University of KwaZulu-Natal, Pietermaritzburg, 2010.

Thesis (PhD)--Stellenbosch University, 2012. / ENGLISH ABSTRACT: This study tested the proposal that fleshy-fruited alien shrubs competed more effectively for avian dispersal services than indigenous shrubs. Several different quantitative measures of increasing complexity were applied to test bird foraging preferences for fruits of two established alien shrubs (Lantana camara, Solanum mauritianum), two emergent alien shrubs (Myoporum tenuifolium, Pittisporum undulatum) and two indigenous shrubs (Chrysanthemoides monilifera, Olea europaea subsp. africana). Overall the simplest quantitative measure, namely that of bird visitation frequency identified foraging preferences of individual bird species for fruits of alien and indigenous shrubs. Moreover, even the more complex measures of numbers of foraging birds, foraging times and products of these in the computed consumption intensity and seed dispersal efficiency were positively correlated to visitation frequency. There was a distinct preference of several, especially heavily fugivorous, bird species for fruits of established alien than indigenous shrubs and several moderately fugivorous bird species preferred fruits of emergent than established alien shrubs. Intricate photographic and experimental approaches were applied to obtain precise numerical data on seed removal rates by birds from the alien and indigenous shrubs. Deficiencies associated with the experimental approach included its inability to discriminate between asynchronous fruit production and fruit ripening, individual foraging bird species, and fruit consumption by other fruit foraging vertebrates. The advantages of the photographic approach were in its provision of a detailed permanent record of individual foraging bird species, their total numbers and precise foraging times and the proportions of whole fruits consumed, as well as other behavioural foraging traits. Tested also was the proposal that frugivorous birds remove larger amounts of fruits from plants with high fruit production and nutritional contents and that the germination of the bird-ingested seeds is enhanced. In compliance with this proposal, canopy fruit mass and monosaccharide content were all positively correlated with the total numbers of seeds removed by birds and viable seeds excreted by birds. Only defecated seeds of the indigenous C. monilifera and alien L. camara displayed enhanced germination following their ingestion by birds Seed dispersal distances of alien and indigenous shrubs by birds were estimated from published records of bird body masses, gut retention times of consumed seed and flight speeds. Estimated seed dispersal distances ranged between 0.41 and 0.81 km for the subset of heavily and moderately frugivorous bird species examined. However, their foraging distances derived from birdring recapture records were much greater, these peaking between 2.5 and 50 km in some heavily frugivorous bird species. It is concluded 1. that bird visitation frequency be applied as the most efficient and practical quantitative measure for gauging bird foraging preferences in future surveys which should involve large complements of indigenous, emergent and established alien species, 2. that flight speeds and gut passage times of ingested alien and indigenous seeds need to be measured in local bird species so long distance seed dispersal potentials can be more precisely determined and 3. that control measures should be focused on eradicating especially emergent alien species with fleshy fruits from urban environments to prevent their transport by birds into adjacent natural areas. / AFRIKAANSE OPSOMMING: Hierdie studie het die voorstel getoets dat uitheemse vlesige-vrug struike meer effektief kompeteer vir voël verspreidingsdienste as inheemse vlesige-vrug struike. Verskeie kwantitatiewe maatstawwe van toenemende kompleksiteit was toegepas om die voedingsvoorkeure van voëls vir vrugte van twee gevestigde uitheemse struike (Lantana camara, Solanum mauritianum), twee ontluikende uitheemse struike (Myoporum tenuifolium, Pittisporum undulatum) en twee inheemse struike (Chrysanthemoides monilifera, Olea europaea subsp. africana) te toets. Algeheel het die eenvoudigste kwantitatiewe maatstaf, naamlik voël besoek frekwensie, voedingsvoorkeure van individuele voël spesies vir vrugte van uitheemse en inheemse struike geïdentifiseer. Addisioneel het die meer komplekse maatstawwe soos voël getalle, voedings tye, asook produkte hiervan in die berekende verbruiks intensiteit en saadverspreiding effektiwiteit almal positiewe korrelasies met besoek frekwensie. Predominant vrugtevretende voël spesies het ‘n duidelike voorkeur getoon vir vrugte van gevestigde uitheemse eerder as inheemse struike. Verskeie meer gematigde vrugtevretende voël spesies het egter vrugte van ontluikende eerder as gevestigde uitheemse struike verkies. Ingewikkelde fotografiese en eksperimentele benaderings was gebruik om presiese numeriese data van saad verwyderingstempo’s deur voëls vanaf die bestudeerde struike te bekom. Tekortkominge geassosieerd met dié eksperimentele benadering sluit in die onvermoë om te onderskei tussen asinkrone vrug produksie en vrug rypwording, individuele kos soekende voël spesies, asook vrug verbruik deur ander vrugvretende werweldiere. Die voordele van die fotografiese benadering was die verskaffing van ‘n gedetailleerde permanente rekord van individuele kos soekende voël spesies, hul totale getalle, presiese voedingstye, proporsie heel vrugte verbruik, asook ander voedingsgedrag eienskappe. Ook getoets was die voorstel dat vrugtevretende voëls groter hoeveelhede vrugte verwyder van plante met hoë vrugproduksie en hoë nutriënt inhoud, en dat ontkieming van voëlingeneemde sade verhoogd is. In nakoming met die voorstel, was blaredak vrugte massa en monosakkaried inhoud albei positief gekorreleer met totale aantal sade verwyder deur voëls en lewensvatbare sade uitgeskei deur voëls. Slegs sade van die inheemse C. monilifera en die uitheemse L. camara het verhoogde ontkieming gewys na inname deur voëls. Saadverspreiding afstande van uitheemse asook inheemse struike deur voëls was bepaal vanaf gepubliseerde rekords van liggaams massa, saad behoud tye in die maag na inname, en vliegspoed. Beraamde saadverspreiding afstande het gestrek tussen 0.41 en 0.81 km vir dié subgroep van predominant en gematigde vrugtevretende voël spesies bestudeer. Nogtans was voedings afstande, afgelei van geringde voël hervangs rekords, veel groter, en het gepiek tussen 2.5 en 50 km vir sommige predominant vrugtevretende spesies. Gevolglik weet ons nou dat 1) voël besoek frekwensie toegepas kan word as die mees effektiewe en praktiese kwantitatiewe maatstaf vir die bepaling van voël voedings voorkeure in toekomstige opnames waar komplementêre groepe inheemse, ontluikende en gevestigde uitheemse spesies bestudeer word; 2) vliegspoed en maag passeer tye van ingeneemde uitheemse en inheemse sade gemeet moet word in plaaslike voël spesies sodat langafstand saadverspreiding potensiaal meer presies bepaal kan word; en 3) beheer maatstawwe moet fokus op die verwydering van spesifiek ontluikende uitheemse vlesige-vrug spesies vanuit stedelike omgewings, om sodoende saad vervoer deur voëls na naasliggende natuurlike areas the verhoed.

South Africa is one of the world's most biologically invaded countries and has spent billions of rands on efforts to eradicate alien invasive plants. Chemical and mechanical control methods have varied in success and the need for integrated management strategies has been realised. This requires a better understanding of all aspects of the invasion process. Some of the most invasive plant species rely on vertebrate dispersers which facilitate long-distance seed dispersal. Frugivory is based on a mutualism in which the frugivores gain a resource and the plants benefit from seed dispersal away from the parent plant. Seed germination itself may either be enhanced, reduced or not affected at all after gut passage. The first aim of this study was to determine if generalist avian frugivores and a fruit bat species (Epomophorus wahlbergi) enhance or decrease seed germination of invasive alien plants in South Africa, by either pulp removal or seed coat abrasion, or if they serve as dispersers only. The second aim was to determine if avian frugivores are able to meet their energetic demands by feeding on a specific alien fruit diet. Finally, we also quantified the nutritional content and morphological characteristics of fleshy fruits of various invasive alien and exotic plant species. Avian frugivores: Red-winged Starlings (Onychognathus morio), Speckled Mousebirds (Colius striatus), and Dark-capped Bulbuls (Pycnonotus tricolor), varied in their effects on the germination success of seeds of four invasive alien species, namely: Lantana camara, Solanum mauritianum, Cinnamomum camphora, and Psidium guajava. However, this was not associated with differences in seed retention times. Similar germination success was observed for avian ingested and de-pulped seeds. This was also observed for fruit bat spat and depulped seeds of Psidium guajava, Melia azedarach, Eriobotrya japonica, and Morus alba. Therefore seed coat abrasion was not important for the germination of these fleshy-fruited invasive alien plants. Pulp removal resulted in significantly earlier seed germination as well as higher seed germination percentages than in the case of whole fruit controls for some of these invasive species. Gut passage is thus important for long-distance dispersal, and in some cases, for enhanced germination of seeds. The invasive Solanum mauritianum and indigenous congener S. giganteum showed similar germination responses, with both ingested and depulped seeds germinating profusely. However, S. giganteum benefited from pulp removal as seeds from whole fruits had less germination. Avian frugivores varied significantly in most energetic parameters calculated when given diets of invasive alien fruit. Speckled Mousebirds and Dark-capped Bulbuls were able to maintain body mass and efficiently process fruits of all four alien invasive plants, while Red-winged Starlings were only able to do so on lipid-rich C. camphora and sugar-rich S. mauritianum. Furthermore, frugivores also adjusted their feeding behavior by eating more nutritionally poor fruit and less energetically rewarding fruit. Fruit bats consumed more fruit per gram body mass than avian frugivores did. They therefore process proportionately more seeds than avian dispersers and thus their role in invasive seed dispersal, which has previously been underestimated particularly in South Africa, is highlighted. Fruits of invasive plant species were similar in morphology, but greater in nutritional content, than fruits of indigenous species. These fruits also contained small, light seeds with approximately only 30% having more than 10 seeds per fruit. The ability of frugivores to efficiently process these fruits and the greater nutritional rewards offered by these provide new insights into why these invasive fruits are preferred by frugivores. In addition, invasive alien plants may have a competitive edge over indigenous species because of their larger reproductive outputs and not necessarily because of greater germination success. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2011.

Complaints of crop damage by cranes on planted maize in the North Eastern Cape, South Africa, have been increasing since the mid-1990‘s, and in some instances severe losses have been reported. Crop damage by the Grey Crowned Crane Balearica regulorum regulorum near the town of Maclear (31º04´S 28º22´E), has been quantified over two growing seasons, and assessed relative to losses caused by foraging Cape Crows Corvus capensis and other feeding damage assumed to be caused by insects. Twelve fields were selected based on previous patterns of crop depredation. Maize seed in seven of the fields was treated with the chemical ‗Gaucho‘ and five fields were planted with untreated maize. In order to determine the source of losses, twenty quadrats (4 m x 4 m) randomly distributed within each field were visited on average every second day, for a period of up to twenty eight days. Results indicate that seed treatments do act as a deterrent to feeding by both cranes and crows, however crane damage is generally insignificant compared to other sources of damage. My study also reviewed past sightings data of the Grey Crowned Crane in an effort to determine if the conversion of former grassland to plantations in this region may have increased foraging activity in maize fields. The data did not allow for clear-cut conclusions regarding changes in distribution or population trends. Conclusions provide direct input into the management of agricultural areas by enabling landowners to take steps to mitigate crop damage. These mitigation measures may either involve the application of seed treatments, or the planting of low risk crops in high risk areas. Future studies should consider the ppossible detrimental effects of chemical seed treatments on crane biology.