This thesis investigated the autecology of the dacine species, Bactrocera cacuminata (Hering) (Diptera: Tephritidae: Dacinae). I specifically focused on the adult phase of the life cycle and resources believed to be significant to this life stage. The prevailing paradigm in dacine ecology predicts that the larval host plant serves as the centre of dacine activity, a state mediated by mutualistic associations with fruit fly-type bacteria. Contrary to predictions, an explicit test of this hypothesis found that the host plant of B. cacuminata, Solanum mauritianum Scopoli, acted almost exclusively as a site for oviposition and larval development. Other key adult behaviours, most notably feeding and mating, were rare at the host plant. Even in disturbed habitats, the paucity of key adult behaviours such as mating was striking. Adult flies of this species were therefore hypothesized to be utilizing other components of their habitat, i.e. resources vital to their life history requirements. Some of the resources that B. cacuminata are known to respond to include sugar, protein, methyl eugenol and the host plant. The latter three resources are believed to be critical in the reproductive success of dacine flies in general. I assessed the physiological status of flies arriving at these resources to determine if flies of different status foraged for resources differently. In dacines, the internal reproductive structures of the male and female flies have been used as predictors of physiological status. I quantified expansion of the male ejaculatory apodeme in B. cacuminata with age of fly and found that there is a threshold apodeme size that is strongly correlated Abstract with sexual maturity. Maturity of female flies could be accurately predicted by ovarian development. Using these methods to assess the physiological and nutritional status of flies arriving at resources (larval host plant, protein and methyl eugenol) in the field, I discovered that only sexually mature and mated females were responding to the host plant, while the males at the host plant were sexually immature. This confirmed the hypothesis that the host plant primarily served as an oviposition site. Additionally, this study revealed that sexually mature males with high nutritional reserves were most commonly collected at methyl eugenol (a plant-derived chemical that elicits a strong response in males of many dacine species) at dusk, the time of peak sexual activity in this species. This indicated that methyl eugenol was perhaps a significant resource in the context of the reproductive behaviour of this species. Methyl eugenol (ME) is one of group of phenyl propanoids to which males of certain species of Dacinae respond. The current hypothesis of the role of these phenyl propanoids is that they function as pheromone precursor chemicals. Response to these chemicals is hypothesized to be a trait under sexual selection. In Bactrocera dorsalis (Hendel), this effect is so strong that a single feeding on ME results in a strong mating advantage up to a month after males feed on the chemical. Bactrocera cacuminata fed on multiple occasions on ME in a laboratory bioassay. After a single 24-hour exposure to ME, investigations of mating competitiveness did not reveal any obvious advantage for ME-fed males over unfed males. However, ME-fed males did enjoy a higher mating success 16 and 32 days after exposure to the chemical, suggesting that some physiological benefits unrelated to the pheromone synthesis was driving this delayed advantage. Investigation of the physiological consequences of feeding on ME revealed no enhancement of nutritional or energetic reserves, suggesting that the delayed mating advantage observed was more likely a chance event. An alternate hypothesis about the proximate function of ME, proposed by Robert Metcalf, is that it serves as a mate rendezvous site. As mating behaviour was notably absent at the host plant, I tested Metcalf’s hypothesis. A field-cage experiment, spatially separating adult resources (host plant, methyl eugenol, sugar and protein) clearly demonstrated that methyl eugenol was functioning as a mate rendezvous stimulus for B. cacuminata. This is the first direct support for Metcalf’s hypothesis. A synthesis of the literature revealed that significantly greater ecological and evolutionary information was required to understand the basis of dacine response to phenyl propanoids. Different dacine species may be utilizing these chemicals differently, even if their evolutionary origin may have been as a plant based kairomone. My studies show that generalizations on the ecology and behaviour of Dacinae, often extrapolated from research on a few pest species, do not hold up in the case of B. cacuminata. This suggests that a more autecological, species-specific approach is required in dacine research, before any predictive generalizations can be made.

Zoological institutions emphasise the importance of excelling in the areas of animal welfare, conservation, education, and research, not only to better the lives of the animals under their care, but to also influence the general population in the pursuit to conserve the natural world. As a result, zoo life is anything but simple. This research project monitored the lives of a captive group of chimpanzees over a two-and-a-half-year period, during which time we explored four research topics while assessing the development of a cognitive research programme and chimpanzee (Pan troglodytes) introductions in a zoo: welfare, cognition, public engagement with science, and animal management. The project’s use of touchscreen technology and on-exhibit research was the first of its kind for the Royal Zoological Society of Scotland’s Edinburgh Zoo. As a result, the researchers placed a great deal of importance not only on assessing the welfare of the chimpanzees throughout training and testing phases, but also assessing the public’s perception of cognitive research being conducted through an internationally broadcast documentary about the project. In the short duration of the project, these research naïve chimpanzees did not fully grasp the concept of video selection in our free-choice activity, but overall, the introduction of a cognitive research programme did not compromise welfare, and the chimpanzees’ repeated interest suggests that chimpanzees found the research to be reinforcing. Partly funded by the BBC, the Chimpcam Project was shown in the UK (broadcast January 2010) and in a variety of other countries, including the United States and Canada (on Animal Planet in 2011). The broadcast allowed us to gather information over the internet on the wider public’s perception of conducting research with great apes in zoos, to complement data collected on visitors to the exhibit itself. Our assessment of the documentary’s impact on public perception showed that it had a positive influence on perceptions of zoo research, scientists, welfare, and the importance of choice for animals. During this research project, a new group of chimpanzees arrived in Edinburgh as part of the international breeding programme for western chimpanzees (Pan troglodytes verus). As the zoo’s focus switched to helping the two chimpanzee groups merge into one, we took the opportunity to apply psychological research to this context, namely the use of video as a research tool and the recognition of the importance of individual differences in response to challenge. The project maintained the cognition and welfare focus by using video introductions (allowing the chimpanzees to watch video footage of the individuals they were about to meet and track the formation of other sub-groups). In addition, personality ratings and chimpanzee behaviour during the visual access period (an animal management technique used prior to physical introductions where the groups could see each other without physical contact) were collected to examine the efficacy of these measures in guiding introductions in order to reduce risk. Personality ratings and behaviours observed during the video introductions could predict the chimpanzees’ behaviour during the physical introductions, however, the visual access period had no predictive power. The welfare implications of the introduction process were also assessed and suggested that: the choice of location (i.e. options of where to be) was more important than the total amount of available space; having individuals removed from your group was more stressful than having individuals added; self-directed behaviour (SDB) performance was context-specific where rubbing significantly increased during periods of uncertainty that were not necessarily negatively valenced; regurgitation and reingestion (R/R) decreased over time; and both in-group members and those of high ranks spent more time grooming others. Overall our data indicate that the chimpanzees coped well with both cognitive challenges and social upheaval during introductions. Despite being regularly studied in captivity and in the wild, chimpanzees have a great deal more to teach us about their world. In order to provide the best welfare for the chimpanzees in our care, we need to understand how research and management practices affect their lives and how the public interpret what we do as researchers. By understanding these aspects of their world, we can better serve those in captivity and influence public opinion on the importance of conserving those in the wild.

Behaviors can exhibit a wide degree of plasticity depending on the environmental context in which they are expressed. Despite this, repeatable differences have been found among and within individuals across a wide range of taxa. For my thesis, I investigated individual differences in foraging and parental care. In the first experiment, I assessed house sparrows (Passer domesticus) for domain-generality among neophobia, habituation and associative learning as they are all responses to novelty. While the results of the study find individual differences in each of these contexts the conclusion supported separate mechanisms for each response (domain-specificity). In the second experiment, I examined how the loudness of brood begging vocalizations influenced parent trip time, food load size brought to the nest and the amount of time spent in the box. The results of this study found individual differences in trip time and the time spent in the box with regard to the initial five seconds of begging loudness during a parent’s visit. Additionally, trip time was also influenced by the change in loudness within a visit. My findings reveal that individual variation may depend on the context in which individuals are measured.

The evolution of elaborate male ornaments via sexual selection is well-understood while the selective pressures acting on female ornaments remains unresolved. Female ornaments in species with strong sexual selection on the male homologue of the ornament were originally thought to result from an intersexual genetic correlation. My thesis explores the evolution of ornaments in females due to direct selection by developing theoretical models and examining the biology of a neotropical mosquito (Sabethes cyaneus) with sexually homologous ornaments coinciding with male-male competition. I began by exploring the morphology of the ornaments in both sexes of S. cyaneus. Sexual dimorphism in the size and shape of the ornaments was slight and both male and female ornaments showed classic hallmarks of sexually selected traits. I then tested for direct selection on S. cyaneus male and female ornaments via mutual mate choice. I found evidence of male, but surprisingly not female, preferences for ornaments. I then further considered the evolution of male mate choice in polygynous species. First, I investigated whether male investment in courtship by S. cyaneus may result in a lower operational sex ratio and thereby reduce the costs associated with male mate choice. Male courtship did pose a significant longevity cost to male S. cyaneus. Second, I explored the possibility that a female preference for male courtship effort may contribute to the benefits of male mate choice in a series of population genetic models. The spread of a male preference gene can be driven by female preferences for male courtship when males court preferred females more. Finally, I found that female S. cyaneus are not benefitting from signalling to increase their mating rate as they are monandrous. My thesis therefore challenges standing sexual selection theory and suggests that sexual selection on females may be more widespread than previously thought. / Felaktigt tryckt som Digital Comprehensive Summaries of Uppsala Dissertations from the Faculty of Science and Technology 729

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Dans cette enquête nous entreprenons d’explorer la notion de comportement animal telle que l’exprime une discipline contemporaine, l’écologie comportementale. Afin de procéder à l’examen d’une notion aussi complexe, positionnée dans un contexte étroit, nous développons et utilisons un outil d’investigation : l’épistémologie historique. De façon générale, cet outil consiste à intégrer en une seule démarche les perspectives d’investigation diachronique et synchronique sur un même thème. Ainsi, pour procéder à l’examen de la notion de comportement animal, nous puisons d’abord dans l’histoire récente de l’écologie comportementale. Nous nous intéressons plus particulièrement à sa filiation avec l’éthologie classique et, après avoir reconstitué la trame historique qui unit les deux disciplines, nous procédons à leur comparaison. Cette seconde étape de notre épistémologie contribue à mettre en évidence plusieurs différences critiques dans la conception du comportement animal qu’endosse chacune des disciplines. Ces distinctions, en faisant ressortir la spécificité de l’écologie comportementale, nous permettent ensuite de nous intéresser à la notion de comportement animal à travers les approches principales que cette discipline mobilise. Ultimement, nous élaborons deux définitions de la notion de comportement animal. La première reflète le statut ontologique du comportement dans la discipline, alors que la seconde correspond à la conception qui se dégage de la pratique des écologues. / In this inquiry I undertake to explore the notion of animal behaviour as it is expounded in a contemporary field of inquiry: behavioural ecology. In order to carry out an analysis of such a complex notion, localized in a very narrow context, I design and use a specific tool of investigation called “historical epistemology”. Simply understood, this tool consists in the integration of diachronic and synchronic perspectives of investigation into a single approach to investigate a circumscribed theme. So, in order to proceed to the analysis of the notion of animal behaviour, I first draw into the recent history of behavioural ecology. I take special interest in its filiation with classical ethology and, after having reconstructed the historical frame that links the two fields together, I proceed to compare them. This comparison, the second step of my epistemology, is used to highlight the characteristics of the animal behaviour conceptions put forward by the two scientific disciplines. These distinctions, bringing to the fore the specificity of behavioural ecology, then allow me to scrutinize the notion of animal behaviour as it is instantiated in the main approaches mobilized by the discipline (especially what I identifies as the “phenotypic adaptationist”, the “phenotypic structural”, the “comparative” and the “by reduction” approaches). Last, I design two definitions of the notion of animal behaviour. The first one reflects the ontological status of the notion in this field of investigation, whereas the second corresponds to the conception underlying behavioural ecologist practices. / Projet réalisé dans le cadre d'une cotutelle avec l'Université de Bourgogne (Dijon, France)

Diet selection and its variation due to difference in forages nutritive value are central processes in plant-herbivore interactions. An incorporating of these parameters may allow predicting diet selection patterns which are a basic underpin for sustainable vegetation management and profitable animal production. To achieve these issues many studies were done two of which were run to study the browse-browser interactions in a sub-humid subtropical savannah, a herd of goats was used as a model browser in natural pasture. Other studies evaluated the nutritive value of five plant species which were the most selected by goats. The objectives of this study were to (i) determine the seasonal patterns of diet selection of goats on woody species and their nutritive value and (ii) to predict the diet selection. The secondary objectivrs were to (i) investigate how plant characteristics such as plant morphology (spinescence, and broad vs. fine leaves), phenology (evergreen vs. deciduous species) and plant chemistry (nitrogen, neutral detergent fibre, acid detergent fibre, acid detergent lignin, condensed tannin, cellulose and hemicellulose) are relate to the diet selection patterns and to (ii) evaluate the nutritive value of the most plant species selected by goats during the different seasons. Two experiments were conducted at Owen Sitole College of Agriculture, KwaZulu-Natal, to determine diet selection of goats grazing/browsing in Zululand Thornveld. The first experiment was done in the early wet season (November 2007) and late wet season (March/April 2008) while the second experiment was conducted in the dry season (June/July 2008), early wet season (November/December 2008) and late wet season (February/March 2009). The relative abundance of browse species in the veld was determined and used to estimate the selection index. In the first part, diet selection was estimated using (i) direct observation of animals (account bouts) by observing every one minute two goats while they were foraging with others for 3 hours in the morning and 1.5 hours in the afternoon on alternate days for a total of six days and (ii) an indirect plant-based method by observing at regular intervals 40 marked branches on ten plants of each browse species every two days for a total of six days. The browse species observed were: Gymnosporia senegalensis, Acacia nilotica, Acacia karroo (Acacia natalitia), Scutia myrtina and Dichrostachys cinerea. In both seasons, the three most selected species according to the plant-based observations were S. myrtina, A. karroo and D. cinerea. Consistently, A. nilotica experienced moderate defoliation and G. senegalensis the least. However, on the basis of the selection index, the species followed the order: A. nilotica > D. cinerea > A. karroo > S. myrtina > G. senegalensis in the early wet season and A. nilotica > S. myrtina > D. cinerea > A. karroo > G. senegalensis in the late wet season. Both methods did not rank species in the same order. The selection index was negatively (P<0.05) correlated to neutral detergent fibre (NDF), acid detergent fibre (ADF) and acid detergent lignin (ADL). In the second part, observations were taken on feeding time of two goats, randomly selected per day for 7-8 days. Observations were made during active foraging periods for 2 hrs in the morning and 1.5 hrs in the afternoon. The duration of each feeding bout and the species of woody plant from which bites were cropped at each feeding station were recorded. Season and plant species affected diet selection and preference. The five main species selected (utilized) by goats in decreasing order were S. myrtina, A. nilotica, D. cinerea, Acacia natalitia and Chromolaena odorata. Scutia myrtina was the most preferred (highest utilisation relative to availability) in the dry season while D. cinerea comprised the greatest proportion in the early and late wet seasons. Scutia myrtina was most preferred in the dry and early wet seasons while A. nilotica was most preferred in the late wet season. Spinescent species were generally selected more than non-spinescent species in all seasons, while fine-leaf and deciduous species were selected more than broad-leaf and evergreen ones in the wet seasons. However, preference for broad-leaf and evergreen species increased in the early wet season. Although plant chemistry varied across seasons, it did not explain the preference of goats for various plant species in this study. Instead, effects of chemistry were species- specific. The nutritive value of the five main species selected by goats was evaluated by chemical composition, in vitro gas production, in vitro degradability and in sacco degradability. Two experiments were conducted for the in vitro studies, the first experiment was to test the effect of season and species, while the second experiment tested the biological effect of tannins using polyethylene glycol 4000 (PEG). The PEG treatment was applied to samples of the early and late wet seasons only. The parameters were maximum gas production (GP), degradation rate (C), lag time (lt), maximum rate of GP at the point of inflection (μ), half time to the maximum gas volume(T1/2), gas produced from fermentation of soluble and slowly degradable fractions (A and B, respectively), their degradation rates (c1 and c2, respectively), apparent (ApDeg) and true degradability (TrDeg), microbial yield (MY), partitioning factor (PF), degradation efficiency factor (DEF) and short chain fatty acids (SCFA). There were wide variations among seasons and species in crude protein (CP), NDF, ADF and condensed tannins. Season and species affected kinetics of gas production, GP, TrDeg, MY, PF and SCFA. Chromolaena odorata had the highest CP (185.8-226.4 g kgˉ¹), GP (87.3-104.1 ml gˉ¹ DM), gas produce from soluble (47.6-50.9 ml gˉ¹ DM) and insoluble (39.8-50.9 ml gˉ¹ DM) fractions during the three seasons compared to other browse species. The TrDeg ranged from 634 to 856 g kgˉ¹ DM. The total SCFA varied between seasons and among species. The addition of PEG decreased TrDeg and PF, and increased GP and total SCFA. The GP, its degradation rate (C) and gas from the soluble fraction were positively correlated to CP without and with PEG. Gas from the soluble fraction was negatively correlated to NDF, ADL and CT; and GP to CT without PEG. In the in sacco study, the parameters of dry matter (DM) and nitrogen (N) degradability were soluble (a) and slowly (b) degradable degradability, the degradation rate (c), potential degradable (PD), effective degradability (ED) and lag time (lt). Season affected PD of DM degradability and ED of both DM and CP. Browse species and its interaction with season affected all parameters except b fraction of CP and interaction did not affected PD of CP. Chromolaena odorata had the highest estimated parameters of degradation during the three seasons compared to other browse species. Based on PD and ED, the plant species followed this decreasing order: C. odorata, A. nilotica, A. natalitia, S. myrtina and D. cinerea. All parameters were positively correlated to CP except the a and b fractions of DM and CP, respectively, and lt of both DM and CP. All parameters were negatively correlated to NDF, ADF and ADL except the b fraction and rate of degradation. Condensed tannins (CT) were negatively correlated with all parameters except the b fraction and PD of DM and CP, and lt of DM degradation. Cellulose was negatively correlated with a of DM and lt of CP degradation. The broad objective of this study was to assess whether plant characteristics, in sacco degradability, in vitro gas production, digestibility or chemical composition (NDF, ADF, ADL, CT and CP) could predict the selection of browse species. The parameters were selection index, plant characteristics (spinescence, or leaves phenology), chemical composition, in vitro gas production, in vitro degradability and in sacco degradability parameters. Most of these parameters were poor for predicting selection by goats of browse species. Spinescence (spn), leaves phenology (phen), NDF, CP and CT accounted for 86% of the variation in selection index (y) = -5.91 - 0.01CT - 0.002NDF + 0.02CP + 6.18spn + 2.43phen; (R2=0.86; n=20; RMSE=0.406; P<0.001; for phen, 1 = evergreen, 0 = deciduous; for spn, 1 = spinescent, 0 = spineless). It was concluded that in vitro gas production, in vitro degradability and in sacco degradability were poor predictors of selection index of browse species used by goats. Spinescence, leaf phenology, condensed tannins, CP and NDF were predictors of diet selection index and suggested that these plant species have potential to be used as feed supplements. Chromolaena odorata has the highest potential as feed protein source in goats. The addition of PEG (tannin binding agent) emphasizes that the inhibitory effect of tannins on rumen microbes was great in tannin-rich feeds. / Thesis (Ph.D.)-University of KwaZulu-Natal, Pietermaritzburg, 2012.

Diet selection and foraging efficiency are primary driving behavioural processes in mammalian herbivore plant interaction. They determine the quality of production and also impact on vegetation, necessitating managerial inputs that will sustain productivity and conserve or improve on its plant species composition. In view of a better understanding of browse/browser interaction in subtropical arid savannah, Nguni goats were adopted as our model browser considering their socioeconomic importance to local, regional and global economy. The primary objective of this research was to determine the seasonal and plant species trend of diet selection indexed by intake and foraging efficiency indexed by intake rate. The secondary objective was to investigate how plant morphological variables such as spinescence, shoot morphology such as long shoots as opposed to short shoots, and broad as against fine leaves, phenological variables such as evergreen versus deciduous species, and plant chemical components including: crude protein, neutral detergent fibre (NDF), acid detergent fibre (ADF), hemicellulose, cellulose, acid detergent lignin and condensed tannins relate and possibly explain the observed patterns. Six browse plants, Acacia nilotica, Acacia robusta, Ehretia rigida, Euclea crispa, Rhus lancea and Ziziphus mucronata were used in the first experiment to determine short term intake during the dry and rainy seasons. In the second experiment, 10 browse plants: Acacia nilotica, Acacia robusta, Combretum epiculatum, Dicrostachys cinerea, Ehretia rigida, Euclea crispa, Gymnosporia buxifolia, Rhus lancea and Ziziphus mucronata were used to determine short term intake rate in the dry, early rainy and late rainy seasons. Diet selection was significantly affected by species (F5,60= 31.87 ; p< 0.05) and the interaction between seasons and species (F,60= 2.52; p= 0.039). Intake was not significantly higher in the rainy relative to the dry season, 3.94 ± 0.42a gDM compared to 3.57 ± 0.42a gDM. Rhus lancea and A. robusta were the most preferred species in both seasons, while Z. mucronata was more preferred in the dry season than E. crispa but in the rainy season it was the opposite. Dichrostachys cinerea and A. nilotica were the least preferred in both seasons, but with improved intake of D. cinerea in the rainy season, whereas that of A. nilotica declined significantly. Intake of plant species with long shoots, which were also broad leafed, was significantly higher than that of old short shoot species (t= -3.99; df= 65.80; p= 0.0002). Similarly, intake of evergreen species was significantly higher than that of deciduous species (t= 5.22; df= 34.02; p< 0.0001). Cellulose in the rainy season positively correlated with intake (r= 0.85; n= 6; p= 0.031), bite number cropped (r= 0.98; n= 6; p= 0.001) and browsing time (r= 0.93; n= 6; p= 0.007). In the dry season there was no correlation of any plant chemical components with intake except cellulose that correlated with the intake variable of bite size (r= 0.87; n= 6; p= 0.023). The following regression equation was generated from the correlation to model intake: Intake= 0.68 + BS (3.07), (R2= 0.71; p< 0.05), where BS= bite size. Intake rate varied amongst species (p< 0.05), as well as seasons (p< 0.05). Instantaneous intake rate in the dry season were highest for A. caffra and G. buxifolia, followed by C. apiculatum, R. lancea, E. crispa, A. robusta and Z. mucronata. The bottom tier comprised of E. rigida, D. cinerea and A. nilotica. At the onset of the rainy season, A. robusta, A. caffra and R. lancea had the highest instantaneous intake rate, followed by C. apiculatum, E. crispa and Z. mucronata. The lowest instantaneous intake rates were derived from D. cinerea, E. rigida, G. buxifolia and A. nilotica. At late rainy season, A. robusta, A. caffra, C. apiculatum, E. crispa and R. lancea yielded the highest intake rate, followed by D. cinerea, G. buxifolia and Z. mucronata in the middle tier, and the least were derived from E. rigida and A. nilotica. Acacia caffra, A. robusta, R. lancea, C. apiculatum and E. crispa featured amongst species with the higest intake rate in all three seasons, whereas G. buxifolia from which one of the highest intake rates was derived in the dry season declined to amongst those with the least in both early and late rainy seasons. Ehretia rigida, D. cinerae and A. nilotica were consistent in all three seasons as the least efficiently foraged Browses with long shoots were more efficiently foraged upon than those with short shoots (t= -3.2; df= 284.64; p< 0.002). Evergreen plant species yielded higher instantaneous intake rate relative to deciduous species (t= 2.02; df= 278.01; p< 0.044). Intake rate of spinescent relative to spineless species was not significantly different but more bites were cropped from spinescent relative to spineless species (t= 2.80; df= 256.77; p= 0.006). Bite sizes were smaller for spinescent relative to spineless species (t= -3.30; df= 227.4; p= 0.001). There were significant positive correlations of cellulose with intake rate (r= 0.7; n= 10; p= 0.023), and with condensed tannins (r= 0.7; n= 10; p= 0.022) in the dry season. Additionally, cellulose also had a positive correlation with bite size in the same season (r= 0.7; n= 10; p= 0.016). At early rainy season, there was no correlation of intake with any of the plant chemical components, whereas, NDF (r= 0.6; n= 10; p= 0.044), hemicellulose (r= 0.8; n= 10; p= 0.012) and condensed tannins (r= 0.7; n= 10; p= 0.032) all correlated positively with browsing time at early rainy season. The following regression equations were derived to model intake rate and some intake rate related variables: (1) intake rate= 0.433 + BSDM (0.081), R2= 0.68, p< 0.05 and BSDM= bite size dry matter; (2) Bite rate= 0.503 –BSDM (0.147), R2= 0.597, p< 0.05 and BSDM= bite size dry matter; (3) Bite size dry matter= 1.713 – BR (4.210), R2= 0.597 ,p< 0.05 and BR= bite rate, and (4) Intake dry matter= 3.804 + BSDM (0.865), R2= 0.607, p< 0.05 and BSDM= bite size dry matter. Diet selection and foraging efficiency were both influenced more by shoot and leaf morphology than by spinescence. Plant chemistry on the other hand had diverse and inconsistent effect on diet selection and foraging efficiency trends. The results therefore have important and far reaching implications for Nguni goat and goat production as a whole in the bush veld of Gauteng and related ecological zones. / Thesis (M.Sc.Agric.)-University of KwaZulu-Natal, Pietermaritzburg, 2012.

Research on aversive control of behaviour has dramatically declined over the past decades. This trend is primarily a consequence of an over-reliance on shock-based procedures, which have been increasingly criticized on ethical, practical, and ecological validity grounds. The continued study of aversive regulation thus requires the development of viable alternatives. Six preliminary experiments, triggered by serendipitous observations of Betta splendens’ reactions to unintended water disturbances, allowed for (a) developing a water flows (WFs) experimental paradigm; (b) confirming the aversive function of WFs; and (c) demonstrating the feasibility of the WFs paradigm as an alternative to the use of electric shock, as it does not involve painful stimulation and carries a higher level of inherent ecological validity. Based on the relevance of free-operant avoidance phenomena (Sidman, 1953a) for the study of aversive control, the fact that these have only been demonstrated in one fish species (goldfish) using shocks, and that the only attempt to show another form of avoidance in Betta splendens produced inconclusive results (Otis & Cerf, 1963), the WFs paradigm was implemented in two experiments aimed at addressing these issues. These studies were aligned with a research program on spatiotemporal analysis of behaviour that has demonstrated, over the course of several decades, that a comprehensive understanding of behavioural processes requires an approach that includes, but is not limited to, the study of rates of discrete responses (e.g., key pecks of a pigeon). Accordingly, a more holistic interpretation of experimental data than is typical for behavioural studies was attained through a combined analysis of the frequency and temporal distribution of a target response (crossings in a shuttle-tank), patterns of swimming trajectories, instances and durations of the aversive stimulus, and the occurrence of behaviour related to different features of the experimental tank. In Experiment 1, Betta splendens exposed to a free-operant avoidance procedure reliably escaped WFs but did not develop avoidance behaviour even though escape improved with practice. Moreover, adding a warning stimulus (curtains of air bubbles - CABs) to the free-operant procedure did not produce increments in avoidance behaviour, as has been demonstrated in other species. Considering these findings, Experiment 2 maintained the same free-operant avoidance contingencies, but escape responses were now scheduled to produce the WFs (punishment and extinction of escape). The result of this manipulation was not a substantial decrease of escape, but an initial large increase of this response, followed by a progressive decrease to approximately pre-punishment levels. In addition, punishment did not result in increased avoidance responding as an alternative response. The explanations for these unexpected findings relate to the duration of the CABs; sign- and goal-tracking effects; uncontrolled stimulation produced by water pump activation/operation; unintended reinforcement (mirror reflections and delay between the pump activation and WFs reaching full strength); and the development of responses that allowed the fish to reduce their exposure to high-intensity WFs (i.e., alternative behaviour). The need for investigating the effects of adjusting the WF procedures to the ecology and biology of Betta splendens is also discussed, particularly in regard to their territoriality and predominant defensive response (immobility) in relation to the experimental apparatuses and the target response (changing compartments).

Bottlenose dolphins (Tursiops spp.) are a widely studied species in marine habitats, however, information on estuarine populations in Australia is very limited. To fully understand the importance of estuaries as habitats for dolphins there needs to be clear quantitative data on dolphin populations and their habitat use in estuaries. This study provides the first published data on Indo-Pacific bottlenose dolphins (Tursiops aduncus) abundance estimates, site fidelity, individual ranging patterns, habitat use, flood impacts and sexual segregation patterns over a 3- year period in the Clarence River (CR) and Richmond River (RR) estuaries in northern New South Wales, Australia.The results indicate that, at present, the CR estuary is sustaining a larger dolphin community with a predominantly resident population compared to the RR estuary, which supports a smaller community with lower site fidelity. The CR estuary dolphin abundance estimate of 71 (62-81 95% CI, CV = 0.07) is more than twice the size of the RR estuary abundance estimate of 34 (19- 49 95% CI, CV = 0.23). Differences in site fidelity were observed between the estuaries with 60% and 37% of identified dolphins determined as residents, 26% and 21% as occasional visitors, and 14% and 42% as transients for the CR and RR, respectively. Resource partitioning was apparent in both estuaries with the mean distance resident dolphins were found upstream from the River mouth being greater than for the occasional visitors and transients.Tursiops aduncus was seen all year round in the CR and RR estuaries, with peak sightings occurring in spring at both sites. In the CR the dolphin population showed consistent seasonal fluctuations, whereas this did not occur in the RR population. In the CR the largest spatial distribution of dolphins in the estuary was observed in spring and winter, while in summer they were primarily restricted to the main estuary channel. Different behaviours that were observed; feeding, socialising, travelling, and milling and resting, were found to be influenced by season, tidal phase and tidal range. In both estuaries the core habitat areas used by the population for feeding consisted of areas with considerable slope near the edge of tidal sand banks, adjacent to deeper channels at the entrances of canals, creeks or artificial breakwalls. In addition, the core habitat areas used by the population for milling and resting behaviour in both estuaries occurred in shallow, sheltered areas, often associated with seagrass beds. Socialising occurred more frequently in the CR throughout most of the deeper waters of the estuary, whereas in the RR it was primarily restricted to a small area of medium depth in the estuary.The major determinant of T. aduncus occupancy in the two estuaries was the flood events that occurred, which resulted in the dolphins abandoning the estuary. The mean predicted probabilities for sighting dolphins during non-flood periods were 0.87 and 0.71, during a flood 0.21 and 0.04, and during a post-flood recovery period 0.83 and 0.80 in the CR and RR, respectively. Principal Components Analysis (PCA) showed that when the dolphins were absent from the estuaries, three components were extracted from the water quality parameters in the CR, and two components in the RR. High loadings from the PCA were associated with the changing salinity, turbidity, pH, dissolved oxygen and temperature associated with the flood events. The return of the dolphins to the estuary following a flood depended on the length and severity of the flood event, but generally the dolphins seemed to prefer waters with salinity levels above 29 ‰. This could be associated with higher salinities being important for their physiological health, or because their prey returned to the estuaries during these higher salinity conditions, more likely a combination of both of these factors.Sexual segregation patterns were observed in T. aduncus populations with mixed gender and female groups mainly recorded in the CR, whereas in the RR female groups dominated the estuary. In the CR, significant differences occurred in sightings between the mixed and female groups in relation to water depth and behaviour. Mixed gender groups were sighted predominantly in deeper water and were involved in social behaviour including sexual behaviour and male herding of females. In contrast, the female groups were observed across all water depths, predominately feeding and also participating in more milling and resting behaviours.The high occurrence of aggressive herding behaviour by males in the CR was significantly different at varying depths, tides and seasons, occurring more often in deeper water, at higher tides and in non-breeding seasons. Female groups were found to utilise the small, shallow tributaries and travel for longer distances up these smaller tributaries than the mixed groups, which were concentrated in the deeper main channel of the estuary.The deeper water of the channels may facilitate the males in herding the females, while the female groups’ habitat selection of shallow estuary areas may provide a sanctuary from aggressive males, access to suitable prey items or prey density for mothers and their calves, or a combination of these factors.This study has provided the first detailed research on T. aduncus dolphin population dynamics, habitat use, occupancy and sexual segregation patterns in two Australian subtropical estuaries. To ensure the long-term survival of both of these dolphin populations, management of future increased anthropogenic disturbances from boat traffic, pollution, dolphin watching, industrial or urban development, over-fishing and habitat degradation of the catchment is needed. Good quality water conditions, the protection of the core feeding areas and small shallow tributaries for females and their calves, and sheltered areas for resting behaviour all need to be maintained for the continued conservation of these important dolphin populations.