京都大学 / 0048 / 新制・課程博士 / 博士(人間・環境学) / 甲第21848号 / 人博第877号 / 新制||人||210(附属図書館) / 2018||人博||877(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 市岡 孝朗, 教授 加藤 眞, 教授 瀬戸口 浩彰, 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

The processes that determine the structure of plant communities are of considerable practical and theoretical interest. Natural enemies such as herbivores, seed predators and pathogens provide one potentially important influence on plant diversity. I investigated the effects of natural enemies on plant diversity in two contrasting, species-rich plant communities (tropical forests in Panama and temperate grasslands in the UK), focusing on pre-dispersal seed predation by insects, and the mortality of seeds and seedlings caused by soil fungi. In Panama I found that pre-dispersal insect seed predators generate significant levels of mortality in multiple tropical tree species, with high heterogeneity in predation rates among individuals and at different forest sites. Insect seed predators were highly host-specific, consistent with a role in enhancing plant diversity. At Upper Seeds, a calcareous grassland site in the UK, I used manipulative experiments to show that soil fungi increase the diversity of plants propagating from soil seed banks. A parallel experiment in Panama, mimicking germination under light gap conditions, revealed differential effects of fungi among sites, with fungicide treatment appearing to increase the diversity of propagated seedlings at some sites but reducing it at others. These results suggest that the influence of soil fungi on pre-emergence mortality can alter plant diversity, even when post-emergence mortality from fungal pathogens is limited. In Panama, I also tested whether enemy-mediated mortality increases with rainfall, potentially contributing to the positive regional correlations widely observed between precipitation and plant diversity. In contrast to predictions, neither pre-dispersal insect seed predation nor the influence of soil fungi on seedling recruitment were affected significantly by site humidity, or (for soil fungi) with experimentally manipulated soil moisture levels. Overall, my results provide evidence that pre-dispersal seed predators and soil fungi can affect plant recruitment and diversity at early life stages, with potential consequences for the community structure of adult plants.

Frugivorous insects, specialised herbivores that consume fruit and seeds, are considered detrimental to host plant fitness. Their direct link to genetic fitness via consumption of plant reproductive tissue, and their negative socioeconomic association with agriculture exacerbates their harmful status. However, empirical testing of insect frugivore effects on host plants, and ecological research on the contribution of insect frugivores to multitrophic frugivory systems, is lacking. In the current study, direct effects of a non-mutualistic, insect frugivore/host plant system were tested and results showed variable effects. Beneficial, detrimental, but predominantly neutral effects on germination and seed production were observed between the Queensland fruit fly (Bactrocera tryoni) and tomato and capsicum plants. Significant effects on seed production were unexpected because infestation occurs after seed set. It was also found that eggplant, although a recorded host of B. tryoni, is inconsistent in its ability to sustain B. tryoni larvae through to its final instar. These results confirmed a simplification and presumption associated with insect frugivore (specifically fruit fly)/host plant interactions. Larval movement, infestation-induced fruit decay, pulp removal and germination were then investigated. For all hosts (tomato, apple and paw paw), treatments infested by B. tryoni decayed significantly quicker and to a greater extent than uninfested treatments, with obvious but variable changes to the texture and appearance. The movement of B. tryoni larvae, pattern of infestation-induced decay and pulp removal was unique and host dependent for all hosts. Only seeds from infested tomato were shown to germinate during the experiment. This indicated that host fruit characteristics are responsible, in part, for variable direct effects on host plant fitness by insect frugivores. Variable direct effects between insect frugivores and host plants, combined with the more rapid decay of infested fruits is likely to have implications for seed dispersal and seed predation by a third trophic level. The characteristics of fruit that are changed by infestation by an insect frugivore were then tested for their effect on a vertebrate frugivore, to illustrate the importance of recognising multitrophic interactions and indirect effects in frugivory. Specifically, seed predating rodents were incorporated into the study and their response to infested and uninfested fruits were recorded, as well as their reaction to the changes in fruit caused by insect frugivores (i.e. texture, smell, larvae presence and sound). Apple and pear infested with B. tryoni larvae were found to attract rodents, while infested tomato and paw paw had a neutral effect on the native rats. This differed from the predominant finding in the literature, which was a deterrent effect on avian seed dispersers. Vertebrate response to fruit infested with insect frugivores therefore, is variable. Assessing the indirect effect of insect frugivores on host plant fitness by attracting or deterring another trophic level requires knowledge of the direct effect between the introduced trophic level and the host plant. For example, the attraction of a seed predator may be as detrimental to host plant fitness as the deterrence of a seed disperser. This illustrates the complexity associated with assessing insect frugivore effects on host plant fitness. Results also indicated that differences in pulp texture, caused by infestation, have a significant effect on rodent preference for infested or uninfested treatments. Pulp texture is likely to effect rodent foraging efficiency, whereas the presence of B. tryoni larvae was observed to be inconsequential to rodent response to fruits. For rodents, and indeed any trophic level motivated by foraging efficiency, this finding raises the issue that for long lived fruiting plants, outside factors such as food abundance and competition for food, may cause a variable response to fruits infested by insect frugivores. From these investigations it has become apparent that insect frugivores are not consistently harmful to host plant fitness, as suggested by their negative stigma, but are likely to contribute variable effects, directly and indirectly, on multiple components of plant fitness and multitrophic frugivory systems.