L’eau et le fourrage sont deux ressources non substituables pour les herbivores dans les écosystèmes arides et semi-arides. La distribution spatiale de l’eau de surface détermine la distribution et l’abondance des espèces dépendantes de l’eau. Cependant les processus impliqués à l’échelle individuelle demeurent méconnus. Treize groupes familiaux d’éléphants d’Afrique (Loxodonta africana) et dix mâles ont été équipé de colliers GPS dans le parc National de Hwange, au Zimbabwe, et à sa périphérie. Les éléphants fourragent autour de multiples points centraux : ils visitent un point d’eau périodiquement toutes les 5h, 24h, 48h ou 72h et s’éloignent plus de l’eau lorsque ils font des trajets de plus longue durée. Pendant la saison sèche, la température augmente et les ressources fourragères s’épuisent à proximité de l’eau. Les groupes familiaux d’éléphants visitent les points d’eau plus souvent en augmentant la fréquence des trajets courts et en abandonnant les trajets de 72h. Cependant, ils parviennent à se rendre plus loin de l’eau pendant les trajets de 24h en augmentant la vitesse de déplacement. Ainsi les patrons de déplacement révèlent que les capacités de locomotion et de navigation des éléphants sont au cœur de leur stratégie d’adaptation à la saison sèche. Malgré cela, ces capacités sont rarement incluses dans les modèles d’approvisionnement dans des environnements hétérogènes. Pendant ces trajets, les groupes familiaux sélectionnent les zones de faible densité de points d’eau à des échelles multiples. La force de la sélection pour ces zones de faible densité augmente avec la longueur du trajet et au cours de la saison. Malgré le fait que l’importance des échelles spatiales soit bien établie dans la littérature, les contraintes associées à l’utilisation de multiples points centraux distribués de manière hétérogène dans le paysage ont été négligé alors que cette distribution détermine le degré d’épuisement des ressources fourragères et les rétroactions sur la sélection de l’habitat. J’ai aussi montré que la faune sauvage évite fortement le bétail et les humains qui les conduisent en périphérie d’une zone protégée pendant la saison des pluies. Cependant cet évitement décline au cours de la saison sèche en raison de l’assèchement des points d’eau et de la raréfaction des ressources fourragères. Les éléphants sont de plus en plus contraints par la distribution de l’eau de surface en saison sèche en raison de l’augmentation de leur besoins en eau tandis qu’ils tentent de maintenir leur approvisionnement en fourrage. Cette étude donne une évaluation quantitative de la contrainte en eau à l’échelle individuelle ainsi que les effets de la distribution en eau dans le paysage sur un grand herbivore. Ces résultats peuvent guider les politiques de gestion de l’eau dans un contexte d’aridification dû au changement climatique. / Water and forage are key non-substitutable resources for herbivores in arid and semi-arid ecosystems. The distribution of surface water determines the distribution and abundance of water dependent animal species yet little is known about the processes involved at the individual level. Thirteen African savanna elephant family groups and ten bulls (Loxodonta Africana) were tracked with GPS collars within and on the outskirts of Hwange National Park, Zimbabwe. Elephants behave as multiple central place foragers: They visit waterholes periodically every 5h, 24h, 48h or 72h and travel further from water during longer trips. During the dry season, temperatures increase and forage becomes depleted closer to water. Elephant family groups visit waterholes more often by increasing the proportion of briefer trips and abandoning 72h trips. However, they forage further during 24h trips by increasing travelling speed. Elephant movement patterns revealed locomotional and navigational abilities are at the core of their coping strategies although they are seldom allowed to vary in most foraging models of animal's use of heterogeneously distributed resources. During these foraging trips, family herds select for areas with low waterhole density at multiple scales. Selection strength for low density areas increases with both distance to water and the advancement of the dry season. Although scaling effects are widely recognized, the effects of the spatial distribution of multiple central places constraining foraging have been ignored yet they determine depletion effects and their feedbacks on habitat selection. I also showed that wildlife strongly avoid livestock and people that herd them at the boundary of a protected area during the rainy season yet avoidance decreases during the dry season when foraging and drinking resources become scares. Elephants are increasingly constrained by surface water availability during the dry season as their drinking requirements increase while they strive to main their forage intake. This study provides quantitative assessment of individual water dependence and of landscape effects of surface water distribution on a large herbivore. These findings can inform surface water management in contexts of aridification resulting from climate change.

京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第21249号 / 理博第4419号 / 新制||理||1634(附属図書館) / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 中野 伸一, 教授 山内 淳, 教授 中務 真人 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

This thesis explains two aspects of animal spatial foraging behaviour arising as a direct consequence of animals' need to drink water: the concentration of animal impacts, and the response of animals to those impacts. In semi-arid rangelands, the foraging range of free-ranging large mammalian herbivores is constrained by the distribution of drinking water during the dry season. Animal impacts become concentrated around these watering sites according to the geometrical relationship between the available foraging area and the distance from water, and the spatial distribution of animal impacts becomes organised along a utilisation gradient termed a "piosphere". During the dry season the temporal distribution of the impacts is determined by the day-to-day foraging behaviour of the animals. The specific conditions under which these spatial foraging processes determine the piosphere pattern have been identified in this thesis. At the core of this investigation are questions about the response of animals to the heterogeneity of their resources. Aspects of spatial foraging are widely commented on whilst explaining the consequences of piosphere phenomena for individual animal intake, population dynamics, feeding strategies and management. Implicated are our notions of optimal foraging, scale in animal response, and resource matching. This thesis addressed each of these. In the specific context of piospheres, the role of energy balance in optimal foraging was also tested. Field experiments for this thesis showed a relationship between goat browsing activity and measures of spatial impact. As a preliminary step to investigating animal response to resource heterogeneity, the spatial pattern of foraging behaviour/impacts was described using spatial statistics. Browsing activity varied daily revealing animal assessment of the spatial heterogeneity of their resources and an energetic basis for foraging decisions. This foraging behaviour was shown to be determined by individual plants rather than at larger scales of plant aggregation. A further experiment investigated the claim that defoliation has limited impact on browser intake rate, suggesting that piospheres may have few consequences for browser intake. This experiment identified a constraining influence of browse characteristics at the small scale on goat foraging by relating animal intake rate to plant bite size and distribution. Computer simulation experiments for this thesis supported these empirical findings by showing that the distribution of spatial impacts was sensitive to the marginal value of forage resources, and identified plant bite size and distribution as the causal factors in limiting animal intake rate in the presence of a piosphere. As a further description of spatial pattern, piospheres were characterised by applying a contemporary ecological theory that ranks resource patches into a spatial hierarchy. Ecosystem dynamics emerge from the interactions between these patches, with piospheres being an emergent property of a natural plant-herbivore system under specific conditions of constrained foraging. The generation of a piosphere was shown to be a function of intake constraints and available foraging area, whilst piosphere extent was shown to be independent of daily energy balance including expenditure on travel costs. A threshold distance for animal foraging range arising from a hypothesised conflict between daily energy intake and expenditure was shown not to exist, whereas evidence for an intermediate distance from water as a focus for accumulated foraging activity was identified. Individual animal foraging efficiency in the computer model was shown to be sensitive to the piosphere, while animal population dynamics were found to be determined in the longer term by dry season key resources near watering points. Time lags were found to operate in the maintenance of the gradient, and the density dependent moderation of the animal population. The latter was a direct result of the inability of animal populations to match the distribution of their resources with the distribution of their foraging behaviour, because of their daily drinking requirements. The result is that animal forage intake was compromised by the low density of dry season forage in the vicinity of a water point. This thesis also proposes that piospheres exert selection pressures on traits to maximise energy gain from the spatial heterogeneity of dry season resources, and that these have played a role in the evolution of large mammalian herbivores.

Doctor of Philosophy / Division of Biology / Anthony Joern / John Blair / Ecological communities are comprised both of species and their interactions. The importance of species interactions is embraced by ecological network analysis, a framework used to identify non-random patterns in species interactions, and the consequences of these patterns for maintaining species diversity. Here, I investigated environmental drivers of the structure of plant-pollinator and plant-herbivore networks. Specifically, I asked: (1) Do global-scale climate gradients shape mutualistic and antagonistic networks? (2) At a landscape scale (within a 3,487 ha research site), how do contrasting regimes of major grassland disturbances - fire frequency and grazing by bison (Bison bison) - shape plant-pollinator network structure? (3) How do fire and grazing affect plant-grasshopper network structure? And, (4) What is the role of plant species diversity in determining plant-herbivore network structure? At the global scale, variability in temperature was the key climatic factor regulating both antagonistic and mutualistic network structural properties. At the landscape scale, fire and grazing had major consequences for plant-pollinator and plant-herbivore communities. In particular, bison grazing increased network complexity and resistance to species loss for both plant-pollinator and plant-herbivore systems. Results from an experimental grassland restoration that manipulated plant diversity suggest that plant diversity directly affects plant-herbivore structure and increases network stability. Collectively, these results suggest that environmental gradients and plant species diversity regulate the network structure of ecological communities. Determining how the structure of ecological interactions change with environmental conditions and species diversity improves our ability to identify vulnerable communities, and to predict responses of biodiversity to global change.

Marine consumers and their prey display plasticity that affects the outcomes of their dynamic interactions as well as community structure and ecosystem function. Aquatic chemical signals induced plasticity in consumers and prey from a broad range of taxonomy (phytoplankton to fishes), sizes (microscopic to macroscopic), and habitats (pelagic to benthic), and this complex plasticity strongly affected consumer-prey interactions. Two fishes,

The effect of redistribution of artificial stock waters from a boredrain to pipes and troughs on the vegetation and soil resources was investigated from December 1994 to October 1998 in a sheep paddock in the semi-arid mulga woodlands of south west Queensland, eastern Australia. The study examined 1) the effect of changing water distribution on the distribution of understorey biomass, sheep, cattle and macropods at the paddock scale; 2) patterns in herbivores, vegetation and soil out from the boredrain and change through time following water redistribution; 3) patterns in herbivores, vegetation and soil out from the new troughs and change through time following water redistribution; and 4) the relative effects of native versus domestic stock on piosphere development out from a trough. Landscape zone and season were major drivers of vegetation and soil dynamics and patterns in this mulga landsystem. At the paddock scale, biomass and woody cover patterns were consistent with historical grazing gradients out from the boredrain, semipermanent waters and previous and current fencelines. There was a high degree of temporal variability in herbivore distribution patterns, not just related to changing water distribution. The native herbivores which presumably have evolved in this system were more likely to be correlated with forage resources than water at the scale of this study. This is in contrast to sheep whose spatial selection of feeding sites was partly influenced by non alimentary factors such as distance to waters, winds and fences, but largely unexplained. While sheep distribution was no longer correlated with distance from the boredrain following piping at the paddock scale, there was no evidence that 1) herbivores were focusing grazing activities around new troughs and 2) that vegetation patterns have changed following water redistribution. The boredrain had distinct gradients in vegetation and soil surface condition associated with it that persisted following rainfall and the lightening and then removal of domestic stock. Gradients in functional plant composition, diversity indices, grass cover and soil stability, revealed zones of reduced production potential parallel to the drain. Following closure of the drain herbivore activity rapidly declined, but there was little evidence of rehabilitation during three years of above average rainfall. Reassessment of the site at 5 yearly intervals over a period in excess of 20 years would better tell the story of change post-drain. Current data suggests the drain induced gradient will continue to persist for many years, providing a ‘ghost of waters past’ In the first two and a half years following changeover from the boredrain to point waters, there was little change to the natural landscape patterns in soil and vegetation out from the southern troughs. Patterns in soil and vegetation around troughs initially reflected patterns of geomorphic zone and tree and shrub distribution. While there was an increase in stocking intensity immediately around the new troughs, there was little evidence of a vegetation and soil response to this increased stocking activity. However increased soil erosion and restricted shrub recruitment close to the troughs perhaps signal the beginnings of change out from the new troughs through time. Comparison between prevailing total grazing pressure, kangaroo only grazing and no large herbivore grazing, found that removal of stock had the effect of dramatically increasing the rate of woody cover change over the period of the study. In contrast the proportion of unpalatable plants increased most at the highest stocking rate with both domestic and native herbivores present. Soil surface condition and plant species also responded to exclosure from domestic stock, indicating that resting paddocks has the potential to improve soil and vegetation condition providing macropod densities are at similar levels to when stock are present. Given that the rehabilitation of degraded areas adjacent to boredrains is unlikely at least in the short term, and that little change has occurred out from new troughs, it is likely that there will be little net change in degraded land as a result of piping bores in mulga landscapes with the level of water availability of this study. Factors likely to influence piosphere development and ecological and management implications for the Boredrain Replacement Program are identified.

Relationships between two taxonomic kingdoms; plants and herbivorous insects, are hypothesized to be a major zone of interaction for generating current day biodiversity; and coevolutionary processes between these intricately linked organisms are hypothesized to maintain diversity in plant secondary chemistry. These metabolites play a key role in plant defence against herbivory and a high degree of intraspecific variation is observed at multiple ecological scales. However, the nature of selection maintaining variation in plant defence profiles is still a major question in evolutionary biology and ecology, and progress towards a deeper understanding is hampered by a lack of studies that take into account ecological context and the multivariate nature of plant defence phenotypes. In this thesis, I employ sophisticated chemical analysis techniques to identify a suite of glucosinolate secondary chemicals, representing different biosynthetic pathways, in the wild cabbage, Brassica oleracea, in natural populations in the UK. I used model-based cluster analysis to explore patterns of association between individual glucosinolates, predicting that as simultaneous resource allocation to multiple defences is likely to be constrained; negative associations between defensive traits should be observed. However, results revealed positive associations between glucosinolates. Therefore co-expression of multiple defences may not be costly for this species. Using this information in conjunction with herbivore surveys and experiments, I show that this mixture has the potential to shape patterns of herbivore abundance and host plant utilization: species-specific responses to variation in glucosinolate phenotypes are discovered at various ecological scales. Thus there is the potential for differential selection on plant chemotypes though species-specific attractions and aversions. By conducting fine scale experiments with herbivore species, I also found that glucosinolate variation has an impact on the counter-adaptations that some brassica specialists have evolved: in order to optimally defend against their own natural enemies, Brevicoryne brassicae aphids sequestering glucosinolates from their host plants must do so selectively, and must choose plants whose chemical profile best matches this behaviour. These findings show that glucosinolate profiles may be under natural selection by herbivores in wild populations, and that reciprocal evolution between these plants and their specialists may continue to promote diversity in secondary metabolites. Together these results highlight the complexity inherent in plant-insect interactions, the importance of field studies and generate a wealth of testable hypotheses for future work.

京都大学 / 0048 / 新制・課程博士 / 博士(人間・環境学) / 甲第20458号 / 人博第808号 / 新制||人||194(附属図書館) / 28||人博||808(吉田南総合図書館) / 京都大学大学院人間・環境学研究科相関環境学専攻 / (主査)教授 加藤 眞, 教授 市岡 孝朗, 准教授 西川 完途 / 学位規則第4条第1項該当 / Doctor of Human and Environmental Studies / Kyoto University / DGAM

Mass mortality events (MMEs) can affect communities through large amounts of carcass biomass and significant reductions in population sizes. I conducted laboratory experiments to compare effects of predator and herbivore MMEs on a community using a study system of convergent lady beetles (Hippodamia convergens), pea aphids (Acyrthosiphon pisum), and fava beans (Vicia faba). I used a factorial design crossing input (addition of carcasses) and mortality (removing 75% of the population) of aphids and lady beetles. Lady beetle mortality reduced the top-down pressure on the aphid population, which increased aphid abundance and decreased plant height. Input of aphid carcasses increased aphid abundance. Input and mortality of aphids caused the aphid population to diminish suggesting that top-down effects of mortality were more significant than bottom-up effects of carcasses. My results show that MMEs can have effects through both bottom-up and top-down processes, which is dependent upon who experiences the MME.

The influence of feeding juvenile female leopard tortoises (Geochelone pardalis, n=18) a commercially available, complete, extruded feed three (3) or seven days (7) per week on dry matter and digestible energy intake, apparent digestibility of dry matter, organic matter, gross energy and fiber fractions, animal body weight and measurements, digesta transit time, rate of passage, and indigestible fill was evaluated. Both feeding frequencies are commonly practiced with captive tortoises. When fed 7 compared to 3 days per week, dry matter and digestible energy intake was greater. Tortoises gained more g BW, but not when adjusted per kg initial BW. When fed 7 compared to 3 days per week, tortoises grew more in plastron width (PW) and carapace height (CH), but not midline straight carapace length (MSCL), and grew more in calculated shell volume (i.e., a calculated estimate of shell volume using MSCL, PW, and CH), with a higher calculated body condition index (BCI). Providing short fasts (i.e., feeding 3 compared to 7 days per week) may be useful in slowing tortoise growth when animals are provided food ad libitum. In general, ad libitum feeding, especially of a highly digestible extruded feed, is not recommended for captive juvenile G. pardalis, especially when offered food daily. With two data points (detected as outliers) removed due to low fecal output (and resulting unrealistically high apparent digestibility of all nutrients analyzed) of two animals when fed 3 days per week, apparent digestibility of cellulose in tortoises fed 7 (n=18) compared to 3 (n=16) days per week was lower, but no differences were detected in DM, OM, GE, or any other fiber fractions analyzed. Transit time (TT1) was shorter and indigestible fill was higher in tortoises (n=18) fed 7 compared to 3 days per week, regardless of percent Cr marker recovered. With four animals removed due to <50% Cr marker recovery, tortoises fed 7 compared to 3 days per week exhibited shorter mean retention time (RGIT), with no differences in digesta transit or indigestible fill. Longer digesta retention when food availability included short periods of fasting may have allowed tortoises to extract more energy from cellulose.