O presente trabalho teve como objetivo avaliar a variação da fertilidade na camada superficial do solo em áreas com cobertura de Mata Atlântica primaria, nos estágios sucessionais iniciais e tardios localizados na Serra do Mar e Serra da Mantiqueira e áreas de pastagem com o objetivo de se quantificar os níveis de degradação e recuperação através de indicadores químicos de impacto ambiental do solo. Foi estudada a disponibilidade de nutrientes nas camadas superficiais do solo em função da ciclagem de matéria orgânica correlacionando com diferentes tipos cobertura vegetal. Foram coletadas amostras considerando cinco pontos por fragmentos distribuídos aleatoriamente coletando o solo nas camadas 0-20 e 20-40 cm de profundidade após ter sido retirado à camada de serapilheira. Foram analisados os atributos químicos relacionados à disponibilidade de nutrientes avaliando-se os teores de MO%, C%, N%, K, Ca, Mg, H, Al, P, e os índices de fertilidade pH, SB, T, V% e m%. Para o estudo da variação comparativa da quantidade da serapilheira foram coletadas cinco amostras aleatoriamente nos fragmentos florestais retirando-se o material orgânico superficial em uma área de 1m2 de pontos previamente escolhidos. A análise dos dados demonstrou que a produção de serapilheira da mata primária foi 41,48% superior à da mata secundária tardia e 62,45% à da mata inicial. O teor de matéria orgânica da camada 0-20 cm da mata primária foi 14,23% superior ao da mata tardia e 25,18% ao da mata inicial. Quanto à disponibilidade de macronutrientes observou-se em geral que na camada 0-40 cm os teores dos macronutrientes se encontram nas classes de alta a média para a cobertura de mata primária e média a baixa para os fragmentos de mata secundárias tardia, inicial e de pastagens. A análise percentual demonstrou que os nutrientes que apresentaram maiores variações da cobertura de mata primária em relação às demais coberturas foi inicialmente o cálcio, seguidos do potássio, nitrogênio e fósforo. O magnésio foi o que apresentou a menor variação. O balanço do estado nutricional do solo demonstrou que a cobertura de mata primária apresenta as melhores condições de disponibilidade de nutrientes para o desenvolvimento das plantas caracterizando como um ecossistema em equilíbrio com o retorno de nutrientes ao solo pelo processo de ciclagem. / In present study we evaluate the fertility variation in superficial layer of the soil in areas with primary Atlantic Forest cover, initial and late successions stages placed in Serra do Mar and Serra da Mantiqueira and pasture areas with aims to quantify the levels of degradation and recuperation through environmental chemical impacts in soil indicators. The nutrients available were studied on superficial layers in function of organic matter cyclage, correlating with different kinds of plant cover. Samples were randomly collected in five points per fragments getting the soil in layers with 0-20 and 20-40 cm of deepness, after litter removal. Chemicals attributes related to nutrients available were analyzed by MO%, C%, N%, K, Ca, Mg, H, Al, P indicators and fertility indexes pH, SB, T, V% and m%. To comparative study of litter variation we collected randomly five samples in forestals fragments, taken off the organic matter in a place with 1m2 from points previously chosen. The analysis showed that litter production from primary forest was 41.48% superior to late secondary forest and 62.45% than initial secondary forest. Organic matter data in layer 0-20 cm from primary forest was 14.23% superior to late forest and 25.18% than initial forest. Macronutrients in 0-40 cm layer is classified high to medium to primary forest cover and medium to low for the fragments of late and initial secondary forest and pastures. The percentage analyses shows that the most nutrients fond in primary forest cover in relationship to others cover was initialing calcium, fallowed to potassium, nitrogen and phosphorus. Magnesium showed less variation. The nutritional balance state of the soil, demonstrate that primary forest cover shows the best conditions of nutrients available to plant growing, characterizing an ecosystem in equilibrium with nutrients returns to soil through cyclage process.

O trabalho foi realizado no Parque Estadual da Serra do Mar - Núcleo Santa Virginia,coordenadas geográficas são: 45 30 a 45 11 O e 23 17 a 23 24 S, com o objetivo de avaliar as diferentes respostas da comunidade de palmeiras em dois escorregamentos ocorridos no verão de 1996 devido a um evento atmosférico (Zona de Convergência do Atlântico Sul). Um dos escorregamentos situado em área de floresta preservada e o outro situado em área de vegetação secundária. Por meio de parcelas circulares de 100m2 alocadas nas bordas das cicatrizes dos escorregamentos e no interior da vegetação do entorno de cada cicatriz, em três cotas altitudinais: base, meio e alto. A comunidade de palmeiras foi representada por cinco espécies em três estádios ontogenéticos (plântulas, jovens e adultos), sendo as espécies: Attalea dúbia, Euterpe edulis, Geonoma gamiova, G. pohliana e G. schottiana. A perturbação causou alterações na comunidade de palmeiras de maneira que as espécies como Euterpe edulis e G. schottiana foram estimuladas pelo aporte de luminosidade provocado pelo deslizamento, enquanto G. gamiova diminuiu drasticamente nestas condições em todos estádios de desenvolvimento. Dentre as quais E. edulis, G. gamiova e G. schottiana apresentaram alteração na estrutura de suas populações com aumento da porcentagem de freqüência dos indivíduos jovens. O padrão de regeneração observado indica que espécies de diferentes estratos florestais respondem de modos diferentes a este tipo de perturbação, entretanto não é possível fazer generalizações para outras áreas.

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The risk of death by predation has been a major driver of group living in many prey animals. Animals must adapt to temporal and spatial variation in predation risk and would benefit from using relevant and reliable sources of information both from conspecifics and heterospecifics to better learn about danger. Research on the effects of group living on antipredator strategy has focused largely on group size. However, sociality is often more complex than simple amalgamations of individuals. Those living in groups are likely exposed to unequal levels of predation risk; some are exposed to more danger than others because of factors related to their age, sex, and spatial or social positioning. An individual’s antipredator strategy should reflect its perceived safety levels. I studied antipredator strategies in blue monkeys (Cercopithecus mitis stuhlmanni) in the Kakamega Forest, Kenya. Blue monkeys are arboreal guenons that live in matrilineally-based social groups and form differentiated social relationships. These social relationships could affect how monkeys respond to variable predation risk. Blue monkeys live in dense, biodiverse rain forests and are preyed upon by both aerial and terrestrial predators. They have a well-developed acoustic communication repertoire and have been known to associate with other primates to reduce predation risk (Cords 1987). I conducted five playback experiments and two sets of observational studies, and used data gathered on social interactions among adult females to further our understanding of how group living affects antipredator strategies. I also used 14 years of social interaction data to explore the heritability of social tendencies. In the first chapter, I present a comprehensive literature review of the connections between group living and antipredator behavior. I describe the effects of group size on antipredator behavior and how research on sociality has shifted towards focusing on individuals’ specific relationships and social connectivity. I then describe several ways in which social connectivity has been shown to influence antipredator behavior. I conclude with future directions and then introduce the dissertation. In the second chapter, I focus on heterospecific eavesdropping. I identified the extent to which blue monkey adult females respond to playbacks of alarm and social calls of two syntopic non-predatory bird species—black-faced rufous warblers (Bathmocercus rufus) and joyful greenbuls (Chlorocichla laetissima). Blue monkeys responded differentially depending on both call type and species. I then evaluated differential responses to conspecific and heterospecific callers, hypothesizing that conspecific signals would trigger stronger anti-predator responses because conspecifics are more relevant signals of risk. I conducted a playback experiment in which adult females were presented simultaneously with one alarm or social call from both conspecifics and warblers (4 combinations of alarm and social calls), or ambient rain forest sound (control). Subjects did not differentiate their responses to simultaneous calls according to the type of playback stimulus. These findings suggest that blue monkeys do not differentiate their responses to alarm calls according to caller relevance. Heterogeneous results among different response variables also highlight the importance of examining multiple modes of antipredator behavior. Next, I examine how an individual’s social connectivity influences its antipredator strategy, hypothesizing that more socially connected individuals would benefit from the proximity of more and closely bonded groupmates in enhancing predator avoidance. In Chapter 3, I evaluate the effects of social connectivity on acute antipredator responses, antipredator vigilance, and responses to signals related to various levels of predator-related threat. I first assessed how social connectivity affects the rate at which adult females exhibit acute antipredator responses (such as diving down in trees, climbing up trees, or alarm calling) and the proportion of responses that are major (lasting >30 s), statistically controlling for age, the presence of an infant, and 2-month “seasons”. I predicted that more socially connected individuals would exhibit less frequent acute antipredator responses because they would be better-informed about risk and therefore would exhibit fewer false alarms. I For the same reasons, I also predicted that they would exhibit more major (vs. minor) responses because false alarms are more likely to involve shorter responses (Cords 1987). Contrary to predictions, however, more closely connected individuals exhibited higher rates of acute antipredator responses, which might reflect their enhanced ability to learn about danger from surrounding groupmates, allowing them to detect more potential threats. There was no evidence that social connectivity was associated with the proportion of responses that lasted >30 s. I also found that the rate of acute antipredator responses and the proportion of responses that lasted >30 s varied with season. I then conducted 90-s focal vigilance follows, to assess how long females exhibit antipredator vigilance after controlling for other social and microhabitat factors (e.g., surrounding vegetation density), which can influence conspecific monitoring and exposure to potential predators. I predicted that more well-connected individuals would exhibit lower levels of antipredator vigilance in the absence of any imminent threats and after controlling for other social and spatial factors. More closely connected individuals who were in the spatial center of their social group did spend less time vigilant, but social connectivity was not associated with vigilance times when subjects were at the group’s edge, where exposure to predators and thus predation risk should be highest and antipredator vigilance should generally be higher. In the spatial center of the group, more closely connected individuals should be in a better position to observe their social partners’ antipredator behavior. Microhabitat also influenced antipredator vigilance in multiple ways, which highlights the spatial variation of perceived predation risk in a complex environment. Lastly, I conducted a playback experiment where I examined responses to signals from conspecifics and heterospecifics that are associated with different levels of threat. I predicted that more poorly connected individuals would respond strongly to all signals that might be associated with predators because they must identify personally whether danger is real, whereas more well-connected individuals would have more differentiated responses because they should be near social partners more frequently and can rely on their partners’ antipredator reactions to assess risk levels. However, although subjects did respond more to direct cues of the predator’s presence (its own calls) than to indirect cues of its presence (alarm calls by conspecifics and heterospecifics), there was no evidence that social connectivity affected responses to playbacks. As expected, stimulus type did affect responses—calls from predators (vs. alarm calls or social calls from non-predators) elicited increased looking responses from subjects, which suggests that stimuli that directly signal predator presence will elicit antipredator behavior regardless of the listener’s social connectivity. Overall, social connectivity seems to play a limited role in blue monkeys’ antipredator strategy but there was some evidence that more well-connected individuals were less vigilant when surrounded by groupmates. The ability to distinguish alarm calls by individual callers has not been well-studied, but animals might benefit from making such distinctions if callers vary in how reliably they signal danger. For decades, researchers have tested whether animals can discriminate callers using the habituation-dishabituation paradigm. After habituating subjects by repeatedly presenting calls of one individual, A, they examine whether subjects dishabituate when they hear the calls of a different individual, B (test stimulus). In Chapter 4, I first review studies that used this paradigm to evaluate whether animals discriminate between conspecific callers and then report on two playback experiments which tested whether wild blue monkeys are capable of such discrimination. My review revealed much methodological variation, particularly in the habituation phase and criteria, statistical analysis, and controls. In experiments, I contrasted two methods of habituation, either presenting a fixed number of stimuli (set after pilot observations) or evaluating responses during the series before progressing to the test. Afterwards, I conducted Wilcoxon signed-rank tests to assess habituation statistically. In the first experiment where I played back a fixed number and rate of calls, it was statistically unclear whether subjects habituated to caller A, despite preliminary observations and similar studies that suggested that the experimental design would be appropriate. Because there was not strong evidence that subjects habituated, I did not evaluate statistically whether subjects differentiated between callers in the full dataset. However, in the second experiment where I assessed habituation during the trial, subjects did habituate to caller A and there was weak support that they dishabituated to caller B, which suggests that caller discrimination may occur. From my experiences, I propose an improved design for studies using the habituation-dishabituation paradigm. Lastly, I explore the mechanisms that drive phenotypic variation in social tendencies (and in turn, social connectivity) in adult females. For natural selection to occur, there must be variation in traits, differentiated fitness benefits based on phenotypes, and heredity or a genetic basis underlying phenotypic variation. The previous chapters highlight the variation in and some of the benefits of social connectivity. In Chapter 5, I conducted an exploratory analysis to examine what factors account for phenotypic variance. Using animal models, I found that both environmental and additive genetic variance accounted for some of the phenotypic variance seen in traits associated with social tendencies (using social connectivity as a proxy). Variance in the social environment (i.e., environmental variance) played a large role in shaping observed phenotypic variation in social connectivity. However, all six of the social network measures examined were weakly heritable, which suggests that there is also a genetic basis for behavioral variation, allowing selection to occur. This dissertation emphasizes the importance of examining both antipredator behavior and sociality using multiple experiments, observations, and measures, while also considering the importance of study species and habitat complexity. The relationship between antipredator behavior and social connectivity is not straightforward and can vary greatly between study systems. Although many of my predictions were not supported, I did find evidence that blue monkeys are receptive to heterospecifics, vary their acute antipredator responses and vigilance based on social relationships with conspecifics, adjust their antipredator vigilance according to spatial positioning, and potentially discriminate between alarm callers. The findings presented here expand our knowledge of how animals learn about predation risk by being attentive to conspecifics and heterospecifics.

京都大学 / 0048 / 新制・課程博士 / 博士(農学) / 甲第21470号 / 農博第2313号 / 新制||農||1064(附属図書館) / 学位論文||H31||N5165(農学部図書室) / 京都大学大学院農学研究科地域環境科学専攻 / (主査)教授 北山 兼弘, 教授 本田 与一, 教授 舟川 晋也 / 学位規則第4条第1項該当 / Doctor of Agricultural Science / Kyoto University / DGAM

Ten species of anoles at Bajo Calima within the Choco of Western Colombia separate into two principal microhabitat groups: forest species, and those inhabiting openings and edges. The ten anoles further separate according to ground and vegetation dwellers. There is a relation at Bajo Calima between the number of anole species and vegetational structural diversity. Anole diversity within a given macrohabitat is by perch microsite/microclimate heterogeneity. These are the two major ecological dimensions along which similarity is limited or resources are partitioned.

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The processes that regulate nutrient access by forest trees exhibit substantial variability on both large and small spatial scales. Explicit study of this spatial variability promotes a better understanding of the structure and function of forests. While the importance of space in ecological processes is being increasingly appreciated, there are major gaps in our knowledge about how space influences plant nutrient supply, particularly within a forest stand. This dissertation consists of three chapters that examine the patterns, drivers, and implications of spatial variation in three main processes that make nutrients available to trees: throughfall nutrient deposition, soil nutrient mineralization, and root system development. In Chapter 1, I use data from a field experiment to examine the effect of fertilization on nutrient transfer from the canopy to the soil via throughfall and litterfall in a tropical rainforest. I demonstrate that at small spatial scales, canopy density controls the flux of nutrients in throughfall, while at large scales, soil fertility is an important control, especially for phosphorus. I also show throughfall can be as important as litterfall in the recycling of certain essential nutrients like potassium, and depending on soil fertility, phosphorus. In Chapter 2, I investigate the small scale spatial patterning in soil nitrogen, a nutrient that frequently limits tree growth, in a temperate forest. By quantifying the degree of spatial inequality and autocorrelation in two plots characterized by different dominant tree species, I show that soil extractable nitrogen pools and net nitrogen mineralization fluxes exhibit a high degree of spatial patterning at scales less than 5 meters, with a majority of nitrogen availability contained within hotspots comprising a small proportion of soil area. I also demonstrate that this spatial patterning affects seedling access to soil nitrogen, which has consequences for seedling growth and survival. Chapter 3 examines how tree species and tree size affect the spatial distributions of root systems in two temperate tree species and explores how differences in root spatial coverage could affect tree nutrient access. I find that the spatial distributions of tree root systems can exhibit dramatic differences between species, with a tradeoff between root spatial coverage and total root length. I also discover that the effect of root spatial coverage on soil nutrient access is highly dependent on the spatial patterning of the soil nutrient, such that tree access to patchy nutrients varies greatly based on tree location within the local soil environment, even for medium-size trees. Together, these chapters characterize important patterns and mechanisms of spatial variation in the processes that regulate tree nutrient access.

One of the primary challenges facing researchers and practitioners in their efforts to address issues of poverty and environment is the need to deepen our understanding of the logic that guides local people's decisions over resource use, particularly among the rural poor whose livelihoods depend on fragile and dynamic environments. This study seeks to identify the set of factors that influences how rainforest people respond to abrupt natural disturbances and resource scarcity through changes in livelihood and resource management practices in two rural poor communities of the Peruvian Amazon. Data were gathered through in-depth survey interviews (n=95 households) between June and December 2003 in the Amerindian community of Arica Viejo (Ucayali River) and the mestizo (ribereno) community of Roca Fuerte (Maranon River). The results reveal that socioeconomic characteristics such as forest experience and knowledge, and access to agricultural land explain striking differences among households in livelihood responses to environmental change, particularly concerning resource use behavior, resilience to disturbance, and the propensity to adopt sustainable resource management strategies.