A novel interpretation of leaf mass variation in plants has been recently proposed based on its relationship with variation in leafing intensity – number of leaves produced per unit remaining above-ground dry mass. It is now widely established that between species variation in these two traits is strongly, negatively (and isometrically) correlated for woody and herbaceous species. Possible fitness advantages of higher leafing intensity (requiring smaller leaf size) can be interpreted in terms of the greater concentration of axillary meristems (larger ‘bud bank’), which may provide greater potential deployment flexibility for growth or reproduction. However, no research to date has addressed how these two traits are related at different scales, specifically at the between-individual (within-species) level, or between sexes in dioecious species. For the majority of 24 herbaceous angiosperms studied here, between-individual (within species) co-variation in leaf size and leafing intensity displayed as an isometric trade-off. For the few species with allometric trade-offs, plants with smaller body size produced disproportionately more leaves, indicating a possible mechanism for promoting reproductive economy when plant size is suppressed (e.g. by competition). For two out of three woody dioecious species, and two out of three herbaceous dioecious species, males tended to produce more, smaller leaves than females, possibly promoting pollen dispersal through a more broadly-spaced floral display (developing from more numerous axillary meristems). In contrast, the larger (but fewer) leaves in females may serve to maximize local photosynthetic resources available for axillary fruit and seed maturation. Males and females did not differ in branching rates associated with differences in leafing intensities, but in one herbaceous species, higher leafing intensity in males was associated with higher flower production than in females. In the larger 24-species data set, there was no evidence that species with higher leafing intensity (providing more axillary meristems) was associated with either a greater magnitude or a greater flexibility in branching or flowering intensity. Additional studies are required to explore the possibility that large ‘bud banks’ of axillary meristems (conferred by high leafing intensity) may promote effective mechanisms for tolerating environmental variability, responding to damage (e.g. from herbivory), and/or maximizing reproductive economy. / Thesis (Master, Biology) -- Queen's University, 2009-10-27 21:20:03.509

No description available.

The Cerrado is a biodiversity hotspot undergoing land conversion with need of broad-extent quantification of species and associated ecosystem function. The effects of species on ecosystems can be tested when functional traits are related to ecosystem properties. The patterns of ecophysiologically-linked leaf traits were characterized with the goals of understanding functional diversity of the above ground biomass for ecosystem characterization and discrimination of this status using remote sensing spectroscopy data. Functional groups of plant life form had more consistently different trait status than taxonomy based groups. Specific leaf area and leaf water content were the most significant traits distinguishing functional groups. Spectral indices from a handheld spectrometer were insufficient to capture the variation of these key traits. Future studies integrating remote sensing to understand the effects of Cerrado functional diversity on ecosystem properties would benefit from incorporating life form functional groups, specific leaf area and leaf water content.

Tropical montane cloud forests (TMCF) are one of the most fascinating, but least understood ecosystems in the world, and the interest in the carbon (C) cycle of TMCFs with regard to carbon sequestration and storage practices has increased rapidly in recent years. One feature that prevails in all TMCFs is a decrease in aboveground net primary productivity (ANPP) and standing biomass and leaf area index (LAI) with increasing altitude, together with the stunted growth form of the trees. This thesis focuses on the input part of the TMCF C-cycle, and investigates the controlling factors on photosynthesis on a leaf, canopy, and ecosystem level in the Kosñipata valley in south east Peru, on the eastern slope of the Andes (13º11’28’’S / 71º35’24’’W). Leaf traits are known to relate to foliar C-exchange, and compared with other altitudinal transect studies of TMCFs, the studied sites had similar altitudinal trends for foliar nitrogen (N) content (though not for phosphorus) and leaf mass per area (LMA), with N content decreasing and LMA increasing with altitude. N concentrations were relatively high and LMA values relatively low, but this observed relationship was consistent with those found in global leaf trait surveys. Examining plant stoichiometry (i.e. N:P ratios), the data suggests that unlike the general hypothesis, the Kosñipata forests are not N limited, except for the study site at 2990 m a.s.l. At the 2990 m a.s.l. site, which is the focal study site of the thesis, photosynthetic parameters Vcmax (the carboxylation efficiency of the Rubisco protein) and Jmax (the electron transport efficiency) proved to be similar to those found in lowland tropical rainforest leaves when expressed on an area basis and standardised to 25 °C (55.6 ± 2.6 and 106.5 ± 5.2 mmol m-2 s-1, for Vcmax and Jmax, respectively). However, when standardised to the mean ambient TMCF temperature of 12.5 °C, both photosynthetic parameters were much lower than ambient tropical rainforest Vcmax and Jmax values. The TMCF Jmax -Vcmax relationships were steeper than found in other tropical biomes, indicating a possible adaptation to the lower light availability in TMCFs because of frequent cloud cover, or a consequence of little atmospheric evaporative demand, which is also due to the humid conditions in this forest type. Although N-Vcmax relationships were significant (P<0.05), the fit was not very strong and the relationship between nitrogen use efficiency (NUE) and Vcmax indicates that TMCF species can be regarded as a different plant functional type compared with other tropical forest types. Diurnal measurements of net photosynthesis (A), stomatal conductance (gs) and leaf water potential (Yleaf) showed that different TMCF species experienced non-contrasting diurnal patterns of Yleaf and gs in the dry season. The observed patterns suggest that some TMCF species can be classified as isohydric species, while others behave anisohydrically. Additionally, in situ gs was not very responsive to these to the range of experienced photosynthetically active radiation (PAR), vapour pressure deficit (VPD) or soil water content (SWC), leading to the conclusion that in the studied TMCF, drought stress does not play a role in C-uptake. When using the measured photosynthetic parameters for up-scaling C-uptake to stand scale with a Soil-Plant-Atmosphere model, simulated annual gross primary productivity (GPP) was 16.24 ±1.6 T C ha-1 yr-1, which is about half the GPP observed in neotropical lowland rainforests. Analyses of the modelled results showed that GPP in this TMCF is mostly controlled by temperature, PAR and leaf area index (LAI) and when increasing these three factors to values found in tropical lowland forest, GPP increased up to 75%. In addition, the modelled results indicate that hydraulic limitations on TMCF C-uptake are very unlikely under current climatic conditions. The modelled results also showed that increases in radiation as a result of less cloud cover do not translate to straightforward increases of GPP. The cloudy conditions of TMCFs, which reduced incident PAR in TMCFs, should therefore not be regarded simply as a negative control on TMCF GPP. Instead, the increase in fraction of diffuse radiation partially offsets the decrease in GPP following the reduction in PAR. Overall, the results of this study show that leaves of Andean TMCF forests have similar C-uptake capacity to tropical lowland rainforests when standardized to similar temperatures, but that for in situ C-uptake temperature, radiation and LAI are the key controls.

Ecologia funcional das plantas trata-se de uma nova abordagem dos dados que associa características morfológicas ou fisiológicas das plantas à sua função no ecossistema. Um único atributo pode estra relacionado a mais de uma função no ecossistema, assim a formação de esquemas de respostas ecológicas, através da associação de mais de um atributo, contribui para a realização de inferências mais seguras. Protocolos têm sido realizados com intuito de padronizar a coleta e análise dos dados, viabilizando a realização de análises globais e previsões mais seguras a respeito da mudança no uso do solo e aquecimento global. Porém, a ocorrência de variações dentro das espécies e em diferentes níveis de escala podem se tornar problemas para tais realizações. Nesse contexto o presente estudo se propõe a avaliar possíveis fontes de variação, bem como avaliar as proporções de variação entre diferentes escalas. As escalas envolvidas foram dentro do indivíduo - foram coletadas dez folhas totalmente expostas nas posições Norte e Sul da copa, que conferem diferentes condições de luminosidade; dentro da mesma espécie ou intrapopulacional - foram coletas cinco indivíduos de cada espécie em cada uma das três subpopulações; e entre espécies - foram coletadas quatro espécies de plantas em cada local. Os atributos avaliados foram: espessura, área foliar específica, conteúdo de massa seca foliar, conteúdos de clorofila a, b e carotenóides, razão clorofila a-b e conteúdo de nitrogênio por massa seca. Os dados foram analisados segundo modelos mistos que verificaram a existência de interações entre as escalas. Em seguida foi realizado uma ANOVA ou um teste de Tukey para verificação se há diferenciação entre os atributos nas diferentes posições da copa. Foram gerados modelos que resultaram em proporções de variação dentro dos indivíduos, entre os indivíduos de toda a espécie, entre as subpopulações e entre as espécies. Também foram realizadas Análises de Componentes Principais para verificação da formação de conjuntos de atributos. Não houve diferença dos atributos foliares entre as duas posições na copa. Somente o conteúdo de nitrogênio apresentou interação na escala Espécie. Houve formação de diferentes esquemas de respostas ecológicas. Schefflera macrocarpa indicou características de espécie de crescimento lento e folhas com longo tempo de vida, Erythorxylum suberosum e Birsonima intermedia apresentaram características mais similares, mas B intermedia apresentou características de folhas de vida mais curta em relação a E suberosum,com investimento especial em clorofila a, que pode estar relacionado à arquitetura de copa da espécie. Symplocos oblongifolia apresentou características de folhas de vida curta. Uma variação marcante e inesperada foi a variação dentro dos indivíduos, que deve ser melhor estudada para averiguação de sua fonte. Estudos de arquitetura de copa, plasticidade fenotípica e que envolvam outras escalas menos detalhadas podem orientar o melhor entendimento das fontes de variação dos atributos foliares paras as espécies desse estudo. / Plant functional ecology is a novel approach that combines morphological or physiological characteristics of plants to the ecosystem function. A single attribute can be related to more than one function in the ecosystem, so the grouping some traits can be taken as ecology strategy schemes contributing to many inferences. Protocols have been undertaken to standardize the acquisition and analysis of leaf traits data, allowing the detection of global patterns that can be used in forecasts and policies regarding land use directions and global warming models. However, the occurrence of variations within the species at different scales would be a source of interference in the data acquisition procedures, affecting the reliability of the interpretation. In this context, the present study aims to evaluate the variation regarding different scales using the proportion of variance of each trait in a group of species. The scales involved were: intra-individual - considering leaves exposed in the North and South canopy parts, which confer different lighting conditions; intra-populational - five individuals of each species in each of the three subpopulations; and interespecific - four plant species at each location. The attributes evaluated were thickness, specific leaf area, leaf dry matter content, content of chlorophyll a, b and carotenoids, chlorophyll ab ratio and nitrogen content by dry weight. Data were analyzed using a mixed models technique to detect the existence of interactions between scales. Then an ANOVA with a Tukey test was performed to check for differences among attributes in different positions of the canopy. We used models encompassing the variations within individuals; among the individuals of all kinds, and among subpopulations among species. We also performed a Principal Component Analysis to explore the relations among sets of attributes at the different scales. We found no difference in leaf traits between the two positions in the canopy. Only the nitrogen content showed interactions at the species scale. The Schefflera macrocarpa showed characteristics of slow-growing species and leaves with long lifetime, and Erythorxylum suberosum and Birsonima intermedia showed similar trends but B.intermedia showed leaves with shorter life spam when compared to E. suberosum , with higher investment in chlorophyll a , which may be related to the canopy architecture of the species. Symplocos oblongifolia showed characteristics of short-lived leaves. We found an unexpectedly high variation within individual level, which should be further studied. Studies of canopy architecture, and phenotypic plasticity involving other scales would contribute to a better understanding of the sources of variation in leaf traits in the species used in this study.

Ecologia funcional das plantas trata-se de uma nova abordagem dos dados que associa características morfológicas ou fisiológicas das plantas à sua função no ecossistema. Um único atributo pode estra relacionado a mais de uma função no ecossistema, assim a formação de esquemas de respostas ecológicas, através da associação de mais de um atributo, contribui para a realização de inferências mais seguras. Protocolos têm sido realizados com intuito de padronizar a coleta e análise dos dados, viabilizando a realização de análises globais e previsões mais seguras a respeito da mudança no uso do solo e aquecimento global. Porém, a ocorrência de variações dentro das espécies e em diferentes níveis de escala podem se tornar problemas para tais realizações. Nesse contexto o presente estudo se propõe a avaliar possíveis fontes de variação, bem como avaliar as proporções de variação entre diferentes escalas. As escalas envolvidas foram dentro do indivíduo - foram coletadas dez folhas totalmente expostas nas posições Norte e Sul da copa, que conferem diferentes condições de luminosidade; dentro da mesma espécie ou intrapopulacional - foram coletas cinco indivíduos de cada espécie em cada uma das três subpopulações; e entre espécies - foram coletadas quatro espécies de plantas em cada local. Os atributos avaliados foram: espessura, área foliar específica, conteúdo de massa seca foliar, conteúdos de clorofila a, b e carotenóides, razão clorofila a-b e conteúdo de nitrogênio por massa seca. Os dados foram analisados segundo modelos mistos que verificaram a existência de interações entre as escalas. Em seguida foi realizado uma ANOVA ou um teste de Tukey para verificação se há diferenciação entre os atributos nas diferentes posições da copa. Foram gerados modelos que resultaram em proporções de variação dentro dos indivíduos, entre os indivíduos de toda a espécie, entre as subpopulações e entre as espécies. Também foram realizadas Análises de Componentes Principais para verificação da formação de conjuntos de atributos. Não houve diferença dos atributos foliares entre as duas posições na copa. Somente o conteúdo de nitrogênio apresentou interação na escala Espécie. Houve formação de diferentes esquemas de respostas ecológicas. Schefflera macrocarpa indicou características de espécie de crescimento lento e folhas com longo tempo de vida, Erythorxylum suberosum e Birsonima intermedia apresentaram características mais similares, mas B intermedia apresentou características de folhas de vida mais curta em relação a E suberosum,com investimento especial em clorofila a, que pode estar relacionado à arquitetura de copa da espécie. Symplocos oblongifolia apresentou características de folhas de vida curta. Uma variação marcante e inesperada foi a variação dentro dos indivíduos, que deve ser melhor estudada para averiguação de sua fonte. Estudos de arquitetura de copa, plasticidade fenotípica e que envolvam outras escalas menos detalhadas podem orientar o melhor entendimento das fontes de variação dos atributos foliares paras as espécies desse estudo. / Plant functional ecology is a novel approach that combines morphological or physiological characteristics of plants to the ecosystem function. A single attribute can be related to more than one function in the ecosystem, so the grouping some traits can be taken as ecology strategy schemes contributing to many inferences. Protocols have been undertaken to standardize the acquisition and analysis of leaf traits data, allowing the detection of global patterns that can be used in forecasts and policies regarding land use directions and global warming models. However, the occurrence of variations within the species at different scales would be a source of interference in the data acquisition procedures, affecting the reliability of the interpretation. In this context, the present study aims to evaluate the variation regarding different scales using the proportion of variance of each trait in a group of species. The scales involved were: intra-individual - considering leaves exposed in the North and South canopy parts, which confer different lighting conditions; intra-populational - five individuals of each species in each of the three subpopulations; and interespecific - four plant species at each location. The attributes evaluated were thickness, specific leaf area, leaf dry matter content, content of chlorophyll a, b and carotenoids, chlorophyll ab ratio and nitrogen content by dry weight. Data were analyzed using a mixed models technique to detect the existence of interactions between scales. Then an ANOVA with a Tukey test was performed to check for differences among attributes in different positions of the canopy. We used models encompassing the variations within individuals; among the individuals of all kinds, and among subpopulations among species. We also performed a Principal Component Analysis to explore the relations among sets of attributes at the different scales. We found no difference in leaf traits between the two positions in the canopy. Only the nitrogen content showed interactions at the species scale. The Schefflera macrocarpa showed characteristics of slow-growing species and leaves with long lifetime, and Erythorxylum suberosum and Birsonima intermedia showed similar trends but B.intermedia showed leaves with shorter life spam when compared to E. suberosum , with higher investment in chlorophyll a , which may be related to the canopy architecture of the species. Symplocos oblongifolia showed characteristics of short-lived leaves. We found an unexpectedly high variation within individual level, which should be further studied. Studies of canopy architecture, and phenotypic plasticity involving other scales would contribute to a better understanding of the sources of variation in leaf traits in the species used in this study.

Orientadores: João Vasconcellos Neto, Gustavo Quevedo Romero / Dissertação (mestrado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-22T08:37:00Z (GMT). No. of bitstreams: 1 Zanatta_MateusFornazari_M.pdf: 1558379 bytes, checksum: 8b7c43c47346a21b0fb6f2df8d0e504c (MD5) Previous issue date: 2013 / Resumo: As aranhas apresentam uma grande variedade de estilos de vida, comportamentos e interações ecológicas, mas pouco se sabe sobre a história natural da maioria delas. Existe uma diversidade de formas de cuidado materno em aranhas e ele costuma ser essencial para a proteção e sobrevivência da prole. A seleção de um local adequado para depositar os ovos é uma das primeiras e mais importantes decisões tomadas pelas fêmeas e pode afetar o desenvolvimento dos ovos, a capacidade de proteção das mães e a facilidade para obter alimento. Aysha piassaguera é uma aranha cursorial da família Anyphaenidae e suas fêmeas constroem ninhos em folhas de plantas da borda da mata, onde depositam seus ovos e cuidam deles até a eclosão e dispersão dos filhotes. Nesse trabalho estudamos alguns aspectos de sua história de vida, mais especificamente seu desenvolvimento (capítulo 1), preferência por locais para construção dos ninhos e oviposição (capítulo 2) e função de seu comportamento parental na sobrevivência dos ovos (capítulo 3). Para o estudo do desenvolvimento, coletamos algumas ootecas e criamos as aranhas em laboratório. As fêmeas depositaram em média 123,7 ovos, que levaram 14 dias para eclodir e mais 9,88 dias para dispersar. As aranhas emergiram já no 2º instar, onde a taxa de mortalidade foi maior (20,51%), cada instar durou em média 27,28 dias, a maturidade sexual foi atingida em torno do 11º instar, após uma média de 250,36 dias após a eclosão, sobrevivendo 77,68 dias como adultas. Investigamos se havia preferência por locais para construção dos ninhos em relação à altura do substrato (próximo ao solo ou à copa) e ao grupo de plantas (Monocotiledôneas ou Dicotiledôneas) comparando a proporção de ninhos construídos nesses lugares, além de analisarmos o quanto algumas características foliares (Razão comprimento/largura, Espessura, Força de dobra e MFA - massa foliar por área) explicam a utilização das folhas para a construção dos ninhos. Os ninhos foram construídos em maior proporção na região próxima ao solo e em Monocotiledôneas, algo possivelmente relacionado ao modo de vida errante e arbustivo desta aranha e ao formato alongado das folhas dessas plantas. Inclusive, a característica foliar que melhor explicou a utilização das folhas foi a Razão comprimento/largura, com folhas mais compridas sendo preferidas, provavelmente por permitirem a construção de um ninho mais simétrico e fechado. Considerando o cuidado materno, testamos a influência da presença da mãe e dos ninhos na sobrevivência dos ovos em três tratamentos: Controle (mãe presente e ninho fechado), Sem Mãe (mãe ausente e ninho fechado) e Aberto (mãe ausente e ninho aberto). A sobrevivência foi maior no Controle do que nos outros dois tratamentos, entre os quais não houve diferença. Como a principal causa de mortes foi ataques de inimigos naturais, isso sugere que a presença da mãe é importante para a proteção dos ovos, enquanto o abrigo sozinho é pouco eficiente nisso. Apesar disso, ele poderia influenciar na manutenção de um microambiente propício ao desenvolvimento dos ovos ou facilitar a defesa dos ovos pela fêmea / Abstract: Spiders display a wide range of lifestyles, behaviors and ecological interactions, but little is known about the natural history of the majority of them. There is a great variety of forms of maternal care in spiders and it is usually essential for the protection and survival of offspring. Selecting an appropriate site for egg laying is one of the first and most important decisions made by females and may affect egg development, the protective ability of mothers and the facility to obtain food. Aysha piassaguera is an anyphaenid hunting spider in which females build box-like nests by folding leaves of forest edge plants, in which females lay their eggs and guard them until hatching and dispersal. In this work we studied some aspects of its life history, particularly, its development (chapter 1), preference for nest building and oviposition sites (Chapter 2) and the effect of parental behavior on egg survival (chapter 3). To study development, we collected egg sacs and reared the spiderlings in the laboratory. Females laid on average 123,7 eggs per egg sac, eggs took 14 days to hatch and another 9,9 days for spiderlings to start dispersing. Spiderlings emerged already in the second instar, which had the higher death rate (20,5%). Each instar lasted 27,3 days, on average, and spiders reached sexual maturity around the 11º instar. From emergence it took an average of 250,4 days for a spider to become adult and the adults survived 77,68 days. We investigated whether females displayed preference for nest construction sites concerning substrate high (near the ground or on plants crowns) and group of plants (monocotyledons or dicotyledons) by comparing the proportion of nests built on those places and we also verified how much some leaf traits (length/width ratio, thickness, folding force and LMA - leaf mass per area) explain the use of leaves for nest construction. Female built their nests mainly near the ground and on monocots, which could be related to this being a wandering species that hunts and takes cover on the lower part of the substrate and to the elongated shape of these plants' leaves, for the leaf trait that best explained leaf use was length/width ratio, with females preferring longer leaves, probably because elongated leaves allow the construction of a more symmetric and sealed nest. Concerning maternal care, we tested whether the presence of the mother and the nest affects egg survival using three treatments: Control (nests intact and mother present), Motherless (nest intact and mother removed) and Opened (nest opened and mother removed). Survival was higher on Control treatment than on the other two, which did not differ between each other. Considering that natural enemies' attacks were the main cause of deaths, this shows how the mother presence is crucial for egg protection, whereas the nest has no role in it. Nevertheless, they may help on the maintenance of a proper microenvironment for egg development or make it easier for the females to defend the eggs / Mestrado / Ecologia / Mestre em Ecologia

This thesis elucidates patterns and drivers of invertebrate herbivory, herbivore diversity, and community-level biomass along elevational and land use gradients at Mt. Kilimanjaro, Tanzania. Chapter I provides background information on the response and predictor variables, study system, and the study design. First, I give an overview of the elevational patterns of species diversity/richness and herbivory published in the literature. The overview illuminates existing debates on elevational patterns of species diversity/richness and herbivory. In connection to these patterns, I also introduce several hypotheses and mechanisms put forward to explain macroecological patterns of species richness. Furthermore, I explain the main variables used to test hypotheses. Finally, I describe the study system and the study design used. Chapter II explores the patterns of invertebrate herbivory and their underlying drivers along extensive elevational and land use gradients on the southern slopes of Mt. Kilimanjaro. I recorded standing leaf herbivory from leaf chewers, leaf miners and gall-inducing insects on 55 study sites located in natural and anthropogenic habitats distributed from 866 to 3060 meters above sea level (m asl) on Mt. Kilimanjaro. Standing leaf herbivory was related to climatic variables [mean annual temperature - (MAT) and mean annual precipitation - (MAP)], net primary productivity (NPP) and plant functional traits (leaf traits) [specific leaf area (SLA), carbon to nitrogen ratio (CN), and nitrogen to phosphorous ratio (NP)]. Results revealed an unimodal pattern of total leaf herbivory along the elevation gradient in natural habitats. Findings also revealed differences in the levels and patterns of herbivory among feeding guilds and between anthropogenic and natural habitats. Changes in NP and CN ratios which were closely linked to NPP were the strongest predictors of leaf herbivory. Our study uncovers the role of leaf nutrient stoichiometry and its linkages to climate in explaining the variation in leaf herbivory along climatic gradients. Chapter III presents patterns and unravels direct and indirect effects of resource (food) abundance (NPP), resource (food) diversity [Functional Dispersion (FDis)], resource quality (SLA, NP, and CN rations), and climate variables (MAT and MAP) on species diversity of phytophagous beetles. Data were collected from 65 study sites located in natural and anthropogenic habitats distributed from 866 to 4550 m asl on the southern slopes of Mt. Kilimanjaro. Sweep net and beating methods were used to collect a total of 3,186 phytophagous beetles representing 21 families and 304 morphospecies. Two groups, weevils (Curculionidae) and leaf beetles (Chrysomelidae) were the largest and most diverse families represented with 898 and 1566 individuals, respectively. Results revealed complex (bimodal) and dissimilar patterns of Chao1-estimated species richness (hereafter referred to as species diversity) along elevation and land use gradients. Results from path analysis showed that temperature and climate-mediated changes in NPP had a significant positive direct and indirect effect on species diversity of phytophagous beetles, respectively. The results also revealed that the effect of NPP (via beetles abundance and diversity of food resources) on species diversity is stronger than that of temperature. Since we found that factors affecting species diversity were intimately linked to climate, I concluded that predicted climatic changes over the coming decades will likely alter the species diversity patterns which we observe today. Chapter IV presents patterns and unravels the direct and indirect effects of climate, NPP and anthropogenic disturbances on species richness and community-level biomass of wild large mammals which represent endothermic organisms and the most important group of vertebrate herbivores. Data were collected from 66 study sites located in natural and anthropogenic habitats distributed from 870 to 4550 m asl on the southern slopes of Mt. Kilimanjaro. Mammals were collected using camera traps and used path analysis to disentangle the direct and indirect effects of climatic variables, NPP, land use, land area, levels of habitat protection and occurrence of domesticated mammals on the patterns of richness and community-level biomass of wild mammals, respectively. Results showed unimodal patterns for species richness and community-level biomass of wild mammals along elevation gradients and that the patterns differed depending on the type of feeding guild. Findings from path analysis showed that net primary productivity and levels of habitat protection had a strong direct effect on species richness and community-level biomass of wild mammals whereas temperature had an insignificant direct effect. Findings show the importance of climate-mediated food resources in determining patterns of species richness of large mammals. While temperature is among key predictors of species richness in several ectotherms, its direct influence in determining species richness of wild mammals was insignificant. Findings show the sensitivity of wild mammals to anthropogenic influences and underscore the importance of protected areas in conserving biodiversity. In conclusion, despite a multitude of data sets on species diversity and ecosystem functions along broad climatic gradients, there is little mechanistic understanding of the underlying causes. Findings obtained in the three studies illustrate their contribution to the scientific debates on the mechanisms underlying patterns of herbivory and diversity along elevation gradients. Results present strong evidence that plant functional traits play a key role in determining invertebrate herbivory and species diversity along elevation gradients and that, their strong interdependence with climate and anthropogenic activities will shape these patterns in future. Additionally, findings from path analysis demonstrated that herbivore diversity, community-level biomass, and herbivory are strongly influenced by climate (either directly or indirectly). Therefore, the predicted climatic changes are expected to dictate ecological patterns, biotic interactions, and energy and nutrient fluxes in terrestrial ecosystems in the coming decades with stronger impacts probably occurring in natural ecosystems. Furthermore, findings demonstrated the significance of land use effects in shaping ecological patterns. As anthropogenic pressure is advancing towards more pristine higher elevations, I advocate conservation measures which are responsive to and incorporate human dimensions to curb the situation. Although our findings emanate from observational studies which have to take several confounding factors into account, we have managed to demonstrate global change responses in real ecosystems and fully established organisms with a wide range of interactions which are unlikely to be captured in artificial experiments. Nonetheless, I recommend additional experimental studies addressing the effect of top-down control by natural enemies on herbivore diversity and invertebrate herbivory in order to deepen our understanding of the mechanisms driving macroecological patterns along elevation gradients. / In dieser Dissertation werden Muster und Determinanten von Herbivorendiversität, von Herbivorieraten durch Invertebraten sowie die Diversität und Gesamtbiomasse von Säugetieren entlang von Höhen- und Landnutzungsgradienten am Kilimandscharo (Tansania) untersucht. Kapitel I liefert Hintergrundinformationen zu den betrachteten Variablen, dem Untersuchungssystem und dem generellen Studiendesign: Zuerst fasse ich den aktuellen Kenntnisstand über die Muster des Artenreichtums und der Herbivorie entlang von Höhengradienten zusammen und erläutere in diesem Zusammenhang verschiedene Hypothesen, die zur Erklärung von Gradienten des Artenreichtum herangezogen werden. Ich erkläutere verschiedene Variablen, die zum Testen dieser Hypothesen erhoben wurden und stelle dar, wie diese den Artenreichtum, die Herbivorieraten und die Biomasse beeinflussen könnten. Anschließend beschreibe ich das Untersuchungssystem, sowie das generelle Design der Studie. In Kapitel II werden die Muster und Determinanten der Invertebratenherbivorie entlang von Höhen- und Landnutzungsgradienten an den südlichen Hängen des Kilimandscharos präsentiert. Auf insgesamt 55 Untersuchungsflächen, die sowohl natürliche als auch anthropogen genutzte Habitate am Kilimandscharo in Höhenlagen zwischen 866 und 3060 Meter über Normalnull (m ü. NN) umfassten, wurden die Herbivorieraten ektophager, minierender und gallbildener Insekten an Blättern erfasst. Die Blattherbivorie war sowohl mit klimatischen Variablen [Jahresmitteltemperatur und mittlere Jahresniederschlagsmenge], der Nettoprimärproduktivität (NPP) und mit funktionellen Blattmerkmalen von Pflanzen [spezifische Blattfläche (SLA), Kohlenstoff (C) / Stickstoff (N)-Verhältnis, sowie N / Phosphor (P)-Verhältnis] assoziiert. Die Gesamtherbivorie zeigte eine unimodale Verteilung über den Höhengradienten, wurde aber sowohl von der Herbivorengilde, als auch vom Habitattyp (natürlich versus anthropogen) beeinflusst. Das C/N-Verhältnis von Blättern war die stärkste Determinante der Blattherbivorie und wurde selbst stark durch die NPP bestimmt. Herbivorieraten sanken mit steigendem C/N-Verhältnis. Das C/N Verhältnis nahm mit steigender NPP zu.- Letztere konnte fast vollständig durch Änderungen der mittleren Jahrestemperatur (MAT) und des Jahresniederschlags (MAP) entlang des Höhengradienten erklärt werden. Damit zeigt unsere Studie, dass sich durch klimatische Faktoren und Energie, welche ihrerseits die Blattchemie beeinflussen und so Variationen in der Blattherbivorie entlang großer Klimagradienten ergeben. In Kapitel III werden die Muster im Artenreichtum phytophager Käfer entlang der Höhen- und Landnutzungsgradienten untersucht und die direkten und indirekten Effekte von klimatischen Faktoren (MAT, MAP), NPP und funktionellen Pflanzenmerkmalen (funktionelle Dispersion, SLA, C/N - und N/P - Verhältnisse) auf diese Muster analysiert. Die entsprechenden Daten wurden auf 65 Untersuchungsflächen, die sowohl natürliche als auch anthropogene Habitate entlang eines Höhengradienten am Kilimandscharo von 866 bis 4550 m ü. NN abdeckten, erhoben. Mittels Kescher wurden insgesamt 3186 phytophage Käfer aus 21 Familien gesammelt und in 304 Morphospezies eingeteilt. Der Artenreichtum phytophager Käfer zeigte eine komplexe, zweigipflige Verteilung entlang der Höhen- und Landnutzungsgradienten. Eine Pfadanalyse ergab, dass sowohl die MAT, als auch NPP positiven direkte bzw. indirekte Effekt auf die Artendiversität phytophager Käfer hatte. Die NPP war positiv mit der funktionellen Dispersion von Blattmerkmalen, ein Maß für die Diversität der Nahrungsressourcen, korreliert. Letztere hatte einen positiven Effekt auf die Diversität der Käfer. Die starken direkten und indirekten Effekte von Klima auf die Diversität und Abundanz von phytophagen Käfern, lassen vermuten dass der Klimawandel in den nächsten Dekaden großen Änderungen der Struktur von phytophagen Käfergemeinschaften bewirken wird. In Kapitel IV untersuchen wir den Effekt von Klima, NPP und anthropogener Störung auf den Artenreichtum und die Gesamtbiomasse von Großwild. Dazu wurden auf 66 Untersuchungsflächen, welche natürliche und anthropogene Habitate in Höhenstufen zwischen 870 und 4550m ü. NN umfassten, Daten zum Artenreichtum un der Abundanz von Großwild mittels Kamerafallen erfasst. Mittels einer Pfadanalyse wurden die direkten und indirekten Effekte von klimatischen Variablen, NPP, Landnutzung, Größe und Schutzstatus der Flächen, sowie der Präsenz von domestizierten Säugetieren auf den Artenreichtum und die Biomasse von Großwild untersucht. Artenreichtum und Gesamtbiomasse dieser endothermen Organismen zeigten eine unimodale Verteilung über den Höhengradienten. Verschiedene Nahrungsgilden zeigten unterschiedliche Muster. Es konnte gezeigt werden, dass NPP und der Schutzstatus der Fläche, aber nicht die Temperatur einen direkten, positiven Einfluss auf den Artenreichtum und die Gesamtbiomasse des Großwildes hatte. Die vom Klima abhängige Nahrungsressourcenverfügbarkeit ist also eine wichtige Determinante im Artenreichtum von Großwild. Die Temperatur hingegen, die den Artenreichtum verschiedener ektothermer Organismen entscheidend prägt, hatte keinen direkten Einfluss auf den Artenreichtum des Großwildes Dafür reagiert das Großwild besonders sensibel auf anthropogene Einflüsse, was wiederum die Wichtigkeit von Schutzgebieten unterstreicht. Obwohl die Muster im Artenreichtum und in Ökosystemfunktionen entlang großer klimatischer Gradienten bereits gut dokumentiert sind, ist das Wissen über die zu Grunde liegenden Prozesse nach wie vor unzureichend. Mit meinen drei Studien über die Muster und Determinanten der Herbivorendiversität, der Herbivorieraten und der Großwildbiomasse trage ich somit zur Verbesserung des mechanistischen Verständnisses solcher makroökologischer Muster bei. Wie die Pfadanalysen zeigten, wurden sowohl der Artenreichtum die Biomasse als auch ökologische Prozesse direkt oder indirekt vom Klima beeinflusst. Es ist somit zu erwarten, dass der vorhergesagte Klimawandel ökologische Muster, biotische Interaktionen, Energie- und Nährstoffkreisläufe in terrestrischen Ökosystemen wesentlich umstrukturieren wird, wobei natürliche Systeme wahrscheinlich besonders sensibel auf den Klimawandel reagieren werden. Meine Ergebnisse demonstrieren auch den Einfluss von Landnutzung auf Artenreichtum und ökologische Prozesse. Da der anthropogene Druck auf die natürlichen Ökosysteme des Kilimandscharos immer weiter zunimmt, sollten objektive Biodiversitätsmaße implementiert werden mit denen man Veränderungen in den Ökosystemen und in Ökosystemldienstleistungen schnell detektieren kann. Meine Ergebnisse basieren auf Beobachtungsdaten, die von bestimmten Nebenfaktoren im Feld beeinflusst werden können. Dennoch ist es mir gelungen mit korrelativen Methoden, Organismen in ihrem biotischen und abiotischen Interaktionsumfeld zu untersuchen – ein Szenario, welches in einem rein experimentellen Aufbau in dieser Form wahrscheinlich nicht geschaffen werden kann. Über weiterführende Experimente könnte jedoch zum Beispiel der Einfluss von Prädatoren auf die Herbivorendiversität und Herbivorieraten quantifiziert werden, welches unser Verständnis über die Determinanten makroökologischer Muster noch vertiefen würde.

The tropical rainforests of the Peruvian Andes are some of the most biodiverse and most vulnerable to climate warming in the world. The Andes are predicted to experience substantial increases in warming of between +2 °C to +5 °C by the end of the century, in addition to an increases in the frequency of high temperature extremes, drought and flood events. The response of these forests to climate change over the next century has global relevance, due to the high levels of endemic species present and the potential role these areas will play as refugia for lowland species. Despite this, the response of tropical montane forests (TMCFs) to climate change remains under-studied. Our current understanding of how Andean species will respond to climate change is based on studies of past compositional changes. Upslope shifts in plant communities of approximately 1.2 - 2.0 m·y-1 have been observed along elevational gradients within Central and South America over the last decade. Based on these migration rates, it has been estimated that the majority of communities will lag behind increases in temperature by 5.5 °C by the end of the century. The implications of this for populations at the trailing range edge is unclear, due to a lack of mechanistic data concerning the acclimatory limits of rainforest species. When faced with rapid warming plant species will need to rapidly adapt, acclimate or migrate in order to survive. In the case of Andean species, migration rates may not be sufficient for a species to remain within its optimal thermal niche and adaptive responses will likely be too slow to be effective, hence individuals will have to acclimate in situ to prevent a decline in performance. The acclimatory ability of species can be quantified by measuring changes in performance, leaf physiology and anatomy in response to experimental manipulations of climate, however such studies are rare within the tropics. Here we carried out a seedling transplant experiment, utilising an extensive 400-3500 m asl elevational gradient in the Peruvian Andes, to simulate climate warming and upslope migration of tree seedlings under real-world conditions. To provide context for the transplant study, natural variation in leaf anatomical traits and physiological stress were explored for twelve species belonging to lowland (LF), mid-elevation (LMF) and tropical montane cloud forest communities. Adults and seedlings from the centre and furthest-most extent of each species' elevational range were studied and compared. Seedlings of each elevational forest community were transplanted downslope and upslope of their local elevational range by the equivalent of ±2 °C and ±4 °C in mean annual temperature. The experiment followed the transplanted seedlings of eleven species over a one year period, monitoring survival, growth and physiological stress (Fv/Fm) of individuals. The acclimatory ability of a subset of these species was quantified by measuring changes in photosynthetic capacity (Vcmax and Jmax), respiratory capacity (Rd) and anatomical traits (Na, Pa, LMA, LDMC) in response to transplantation. The results showed that within the natural population there was little evidence of leaf trait acclimation to elevational shifts in climate, but also little evidence of physiological stress at the trailing range edge. There were however differences in the leaf trait strategies employed by each elevational community, increasing in abiotic stress-tolerance with elevation. Physiological stress was greatest in the seedling population and, unlike the adult population, increased slightly at the trailing edge. This indicated that seedlings were more vulnerable to warming than their adult counterparts and at mid-elevations TMCF seedlings were more vulnerable than LMF seedlings. Seedling survival and growth declined in response to transplantation away from the home elevation for the majority of species, with upslope declines as a result of abiotic limitations, and downslope declines due to biotic limitations. All seedlings were found to be able to acclimate their respiratory capacity in response to transplantation, however this was not the case for photosynthetic capacity. LMF species performed significantly better than TMCF seedlings with transplantation, demonstrating a greater acclimatory capacity for photosynthesis. LMF species were able to adjust Jmax in order to maintain rates at ambient temperatures, but were not able to upregulate Vcmax upslope, whereas TMCF species were not able to respond in either transplant direction. Overall, these findings suggest that under moderate warming scenarios LMF species will have a competitive advantage over TMCF species at mid-elevations, gradually expanding their range into TMCF species' habitat over the next century. As a result of this and due to the slow pace of upslope migration, we predict that TMCF species will undergo range retractions and possible extinctions. The speed of this response will be determined by the trajectory of future warming and the frequency of extreme climatic events.

Transições de floresta-campo são encontradas em diversas regiões do mundo. Sob condições climáticas favoráveis, tem-se observado um padrão de aumento na densidade de lenhosas e expansão florestal sobre áreas de vegetação campestre. Este trabalho tem como objetivo identificar diferenças de composição de espécies lenhosas e composição funcional entre comunidades florestais e de transição, bem como identificar padrões funcionais destas comunidades em resposta a gradientes de clima e solo. Para tanto, foram coletados dados em 18 áreas de transição floresta-campo no sul do Brasil, considerando a densidade de espécies lenhosas e atributos foliares mensurados em cada habitat (floresta e transição). Os sítios de amostragem foram descritos por variáveis de clima e solo, gerando assim três matrizes ambientais (E): tipo de habitat, climática e edáfica. A análise dos dados envolveu ajustes de Procrustes entre matriz T (atributos médios da comunidade ponderados pela abundância das espécies) e matriz E (r (TE)) para detectar padrões de convergência de atributos, e entre diversidade funcional (R) e matriz E (r (RE)) para detectar padrões de divergência, relacionando estes padrões a cada matriz E. Os resultados indicaram padrões de convergência e divergência em relação à matriz de habitat. Comunidades florestais e de transição diferiram em termos de média de SLA e área foliar, e também quanto à diversidade funcional (ambos com valores maiores na floresta). Considerando os gradientes ambientais, as comunidades de ambos os habitats apresentaram padrões de convergência com o clima e o solo. Em matéria de clima, o principal resultado foi em relação ao SLA, com valores mais altos em áreas de florestas estacionais. Quanto ao solo, as comunidades florestais demonstraram uma associação de SLA e espessura da folha com o gradiente de matéria orgânica / fertilidade, porém as comunidades de transição não apresentaram padrões claros. Padrões de divergência em relação ao solo foram observados para ambos os habitats, mas só a floresta apresentou divergência em relação ao gradiente climático. Concluímos que, apesar das diferenças locais entre habitats em termos de composição de espécies lenhosas e estratégias funcionais, as comunidades de transição e de floresta estão respondendo de forma semelhante aos gradientes climáticos regionais. No geral, as espécies lenhosas demonstram ter estratégias funcionais relacionadas a atributos foliares que tem possibilitado o processo de adensamento de lenhosas em ecossistemas campestres em áreas de transição de floresta-campo. / Forest-grassland transitions are found in many different regions of the world. Through favorable climatic conditions, a pattern of woody encroachment and forest expansion over open grassy areas is observed. This work aims at identifying species composition and functional differences between forest and transition communities concerning woody plants, as well as functional patterns of communities in response to climate and soil gradients. We collected data in 18 forest-grassland transition areas in southern Brazil, considering woody plant species density and leaf traits that were measured for each habitat (forest and transition). Sites were described by climate and soil variables, leading to three different environmental matrices (E): habitat-type, climatic, and soil gradient. Data analysis involved Procrustes adjustment between matrix T (community-weighted mean traits) and matrix E (r(TE)) to detect trait-convergence, and between functional diversity (R) and matrix E (r(RE)) to detect patterns of divergence related to each matrix E. Results showed convergence and also divergence concerning the habitat-type matrix. Forest and transitional communities differed in terms of SLA and leaf area community-weighted means, and also in functional diversity. Concerning the environmental gradients, either forest or transition habitats presented convergence patterns with climate and soil gradients. Main results concerning climate were related to higher SLA at seasonal forest sites. As for soil, forest communities have demonstrated an association of SLA and leaf thickness with the organic matter/fertility gradient, but transition didn’t present clear patterns. Concerning alpha-divergence both habitats responded to soil gradients, but only forest presented divergence concerning climate. We conclude that although woody species of both habitats locally differed in species composition and their functional strategies, communities of forest and grassland transitions are responding similarly to broader climate gradients. Overall woody species seem to have leaf traits strategies that enabled the encroachment process of grassy ecosystems in forest-grassland transitions.