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京都大学 / 0048 / 新制・課程博士 / 博士(理学) / 甲第19529号 / 理博第4189号 / 新制||理||1601(附属図書館) / 32565 / 京都大学大学院理学研究科生物科学専攻 / (主査)教授 大串 隆之, 教授 中野 伸一, 教授 曽田 貞滋 / 学位規則第4条第1項該当 / Doctor of Science / Kyoto University / DGAM

Thesis (PhD)--Stellenbosch University, 2015. / ENGLISH ABSTRACT: Understanding how divergent selection results in the evolution of reproductive isolation (i.e. speciation) is an important goal in evolutionary biology. Populations of herbivorous insects using different host plant species can experience divergent selection from multiple selective pressures which can rapidly lead to speciation. Restio leafhoppers are a group of herbivorous insect species occurring within the Cape Floristic Region (CFR) of South Africa. They are specialised on different plant species in the Restionaceae family. Throughout my thesis I investigated how bottom-up (i.e. plant chemistry/morphology of host plant species) and top-down (i.e. predation and competition) factors drive specialisation and divergence in restio leafhoppers. I also investigated interspecific competition as an important determinant of restio leafhopper community structure. In chapter 2 I quantified host specificity of restio leafhopper species within a local community for 24 months. I found that restio leafhopper species are highly host specific and potentially synchronised with the growth phases of their host plants. In chapter 3 I used a network metric, modularity, to determine whether host plant partitioning in a restio leafhopper community is non-random (i.e. driven by a deterministic process). This metric allows the identification of the components underlying host plant partitioning (modules). I then performed experiments to show that modules, and therefore host plant partitioning, can mostly be explained by preference and performance relationships (i.e. bottom-up factors). In chapter 4 I used null models to test whether niche partitioning in restio leafhopper communities is a general pattern across the landscape. I found non-random niche partitioning, which results from strong host specificity, in all investigated restio leafhopper communities. In addition, I performed binary host choice experiments in the presence and absence of interspecific competition, but found no evidence that interspecific competition narrows host preferences. These findings suggest that host specificity, the cause of niche partitioning, is likely shaped over evolutionary time. Sampling multiple interaction networks across the CFR, in chapter 5, I tested whether restio leafhopper populations are more host specific in species rich communities and regions in the CFR than in species poor communities and regions. I found no positive relationship between restio leafhopper species richness and host specificity at any scale. These findings suggest that specialisation is not driven by interspecific competition. In chapter 6 I investigated host shifts in Cephalelus uncinatus. C. uncinatus has a broader distribution than any single restio species that it can use; suggesting that host plant related divergence may result from geographic range expansion. I found that allopatric and parapatric populations, but not sympatric individuals, using different host plants have divergent host preferences. I also found evidence for morphological divergence in traits related to predator avoidance in population pairs that exhibit divergent host preferences. My findings emphasise the importance of both bottom-up and top-down factors, with the exception of interspecific competition, as determinants of specialisation and divergence in restio leafhoppers. I find no evidence that interspecific competition is an important force structuring restio leafhopper communities. Instead, strongly niche partitioned community structure appears to emerge from the speciation process. / AFRIKAANSE OPSOMMING: Die wyse waarop uiteenlopende seleksie lei tot die evolusie van seksuele isolasie (n.l. spesiasie) is ‘n belangrike vraag in evolutionêre biologie. Plantetende insekpopulasieses wat verskillende gasheerplante gebruik kan onder uinteenlopende veelvoudige seleksie wees en vinnig spesiasie ondergaan. Restio-blaarspringers is ‘n groep plantetende insekspesies wat gespesialiseerd is op verskillende plantspesies in die restio familie. In my tesis ondersoek ek die onder-op (n.l. plantchemie en morfologie) en bo-af seleksiekragte (n.l. predasie en kompetisie) wat lei tot gasheerspesialisasie en -spesiasie in restio-blaarspringers. Ek ondersoek ook die belangrikheid van tussen-spesieskompetisie in gemeenskapsorganisasie. In hoofstuk 2 het ek gasheerspesialisasie gekwantifiseer in ‘n klein restio-blaarspringergemeenskap oor 24 maande. Ek het gevind dat restio-blaarspringers hoogs gasheerspesifiek is en moontlik met die groeifase van hul hoofgasheerplante gesinchroniseerd is. Ek het in hoofstuk 3 ‘n netwerkmetriek, modulariteit, gebruik om te bepaal of restio-blaarspringers se gasheerverdeling nie-stokasties is (n.l. deur deterministiese prosesse veroorsaak is). Hierdie metriek laat ‘n mens toe om die komponente van gasheerverdeling (modules) te identifiseer. Deur middel van eksperimente het ek bepaal dat modules, en dus gasheerverdeling, deur gasheervoorkeur en prestasie (onder-op prossesse) verduidelik kan word. In hoofstuk 4 het ek ondersoek of gasheerverdeling algemeen is. Ek het deur middel van nulmodelle gewys dat gasheerverdeling algemeen is en veroorsaak is deur sterk gasheerspesifiekheid. Ek het ook voorkeureksperimente uitgevoer in die teenwoordigheid en afwesigheid van tussen-spesies kompetisie. Hier het ek geen teken gevind dat huidige tussen-spesies kompetisie gasheervoorkeur beïnvloed nie. My bevindinge in hierdie hoofstuk stel dus voor dat spesialisasie, die oorsaak van gasheerverdeling, oor evolutionêre tyd gevorm word. In hoofstuk 5 het ek ondersoek of populasies van restio-blaarspringers meer gasheerspesifiek is in restio-blaarspringerspesies ryke gemeenskappe en streke as populasies in spesies-arm gemeenskappe en streke. Ek het geen positiewe korrelasie tussen spesiesrykheid en gasheerspesialisasie gevind nie. Dit dui daarop aan dat gasheerspesialisasie, en dus gasheerverdeling, nie deur tussen-spesies kompetisie veroorsaak word nie. In hoofstuk 6 het ek gasheerplantgekoppelde divergensie ondersoek in Cephalelus uncinatus. C. uncinatus se verspreiding is breër as enige restio-spesies wat dit kan gebruik. Dit stel voor dat verspreidingvergroting gasheerverskuiwing mag veroorsaak. Ek het gevind dat populasies wat verskillende plante in allopatrie en parapatrie gebruik uiteenlopende gasheerkeuses maak, maar insekte wat verskillende plante in sympatrie gebruik wys nie ontwrigtende gasheerkeuses nie. Die populasies wat uiteenlopende gasheerkeuses getoon het, het ook verskillende morfologiese teen-predasie eienskappe getoon. Dit dui daarop aan dat predasie belangrik mag wees vir spesiasie in restio-blaarspringers. Die bevindinge van my tesis dui daarop aan dat beide onder-op en bo-af seleksie belangrik is vir gasheer-spesialisasie in divergensie. Nietemin, tussen-spesies kompetisie is nie ‘n belangrike bron vir spesialisasie, divergensie of gemeenskapsorganisasie nie. Gemeenskapsorganisasie is klaarblyklik slegs ‘n gevolg van die spesiasie proses.

Herbivory, fact of animals feeding all, exudations, tissues or parts of plants, is pointed like one of the most important points in the vegetation structuralization and diversity. That can be influenced by some factors, the most importants are seasonality, spatial variation, environment and plant-herbivory-predator interactions. Native species from Cerrado are also influenced by fire. In response to herbivory attack, the plants have different defenses strategies, it can be chemical, physical and/or biotic. The aim of this study was analyze the impact of herbivory action in Chamaecrista neesiana (Caesalpinoidea), in pre and post natural burn periods, trying to understand the phenologic variation importance to the plant in response to herbivory and fire damage. The study was conducted in Clube Caça e Pesca Itororó de Uberlândia, Uberlândia-MG, in two stages. Firstly, pre-fire (June to September, 2004) and the second, post-fire (November, 2004 to February, 2005). The Burn occurred accidentally in the study reserve in October, 2004. For the study was marked 46 plants of Chamaecrista neesiana (Caesalpinoidea) shared in two groups: the Control one (n=21; without herbivorous insects removal) and the Treatment one (n=25; with herbivorous insects removal). On the plants with herbivorous removal was sprayed weekly a non-nitrogen and non-phophorus based insecticide. This kind of insecticide was used to do not have plants unnatural fertilization. The application of the insecticide occurs by covering the plants with a plastic during five minutes for, more than neighbor plants protection, increases the product effectiveness. The removal of herbivorous insects in Treatment Group occurs also by manual removal. The insects were also collected in non-marked plants, by the beating method and manual collection. The herbivory was measured monthly by visual measure, using a Grid for comparison by the virtual sharing, in eight parts, on the leaf of C. neesiana. Monthly was also observed the phenology dates, the vertical and lateral growing. On the young leafs was observed a lot of oil gland, to verify their function was get on 20 control pitfalls with water and detergent and others 20 was dirty on the edge of the pitfalls with this oil. The herbivory tax was smaller in the protected against herbivorous insects group than the non-protected one, and after the burn. We suggest that this fact occur because of the higher production of young leafs after burn, that happens because in the young leafs have the oil gland production, and we found that this oil gland works like a natural defense by insects, by the collection of more abundance in pitfalls without oil. On this way, was verified more susceptibility to foliar herbivory in mature stages. In C. neesiana, was not found compensatory effect of herbivory, because the vertical and lateral growing do not differ in the both groups (with and without herbivorous insect removal). The phenology showed the boom during the dry season (pre-fire), in the same period of the biggest diversity of insects, and in response to fire the flowering was anticipate in the years 2003 and 2005 (after burning) and occurred gradually, differently the year 2005 (before burning) that the flowering occurred in boom. / A herbivoria, fato de animais se alimentarem do todo, de exsudações, de tecidos ou partes de plantas, é apontada como um ponto importante na estruturação e diversidade da vegetação. Esta pode ser influenciada por diversos fatores dentre eles destacam-se a sazonalidade, a variação espacial, o ambiente e as interações planta-herbívoro-predador. Espécies nativas do Cerrado são, também, influenciadas pelo fogo. Como resposta ao ataque de herbívoros, as plantas apresentam diferentes estratégias de defesa, podendo ser químicas, físicas e/ou bióticas. O objetivo principal desse estudo foi avaliar o impacto da ação de herbívoros em Chamaecrista neesiana (Caesalpinoidea), nos períodos pré e pós-queimada natural, buscando compreender o papel da variação fenológica da planta em resposta à herbivoria e à ação do fogo. O estudo foi realizado no Clube Caça e Pesca Itororó de Uberlândia, Uberlândia-MG, em duas etapas. A primeira, pré-fogo (Junho a Setembro de 2004) e a segunda, pós-fogo (Novembro de 2004 a Fevereiro de 2005). A queimada ocorreu de forma acidental na reserva de estudo no mês de outubro de 2004. Foram marcadas para o estudo 46 plantas de Chamaecrista neesiana (Caesalpinoidea) divididas em dois grupos: um Grupo Controle (n=21; sem remoção de herbívoros) e o outro Grupo Tratamento (n=25; com remoção de herbívoros). A remoção de herbívoros foi feita pela aplicação semanal de inseticida, não contendendo compostos à base de nitrogênio ou fósforo para que não houvesse adubação involuntária das plantas. A aplicação do inseticida se deu com o ensacamento das plantas durante cinco minutos para, além de proteger plantas vizinhas, aumentar a eficácia do produto. Houve também a remoção manual dos insetos no Grupo Tratamento. Insetos também foram coletados em plantas não marcadas, através dos procedimentos de batimento e coleta manual. Foi feita a medida visual mensal da herbivoria, utilizando-se um Grid de comparação pela divisão virtual em oito pedaços dos folíolos de C. neesiana. Mensalmente observaram-se, também, os dados fenológicos, bem como os crescimentos vertical e lateral. Foi observada a presença de glândulas de óleo em folhas jovens, para verificar sua função foram montados 20 armadilhas contendo somente água e detergente em seu interior e outras 20 foram lambuzadas nas bordas com o óleo de folhas jovens. A taxa de herbivoria foi menor no grupo protegido contra ação de herbívoros bem como após a queima, sugerimos que este fato se deve à maior produção de folhas jovens após a queimada, pois em folhas jovens há a produção de glândulas de óleo, as quais constatou-se servirem como defesa aos insetos, pela coleta de maior abundância em armadilhas sem óleo. Desta forma, constatou-se que há maior suseptibilidade a herbivoria foliar nos estágios mais maduros. Não houve efeito compensatório da herbivoria em C. neesiana, pois seu crescimento vertical e horizontal não diferiu nos grupos (remoção ou não de herbívoros). Em relação à fenologia C. neesiana, apresentou seu pico de floração durante a estação seca (pré-fogo), coincidindo com a maior diversidade de insetos, como resposta ao fogo houve antecipação da floração após a queima nos anod de 2003 e 2005 (depois do fogo) e oco / Mestre em Ecologia e Conservação de Recursos Naturais