Investigation of Fundamental Processes Governing Secondary Organic Aerosol Formation in Laboratory Chambers

Our understanding of the processes and mechanisms by which secondary organic aerosol (SOA) is formed is derived from laboratory chamber studies. In the atmosphere, SOA formation is primarily driven by progressive photooxidation of SOA precursors, coupled with their gas-particle partitioning. In the chamber environment, SOA-forming vapors undergo multiple chemical and physical processes that involve production and removal via gas-phase reactions; partitioning onto suspended particles vs. particles deposited on the chamber wall; and direct deposition on the chamber wall. The main focus of this dissertation is to characterize the interactions of organic vapors with suspended particles and the chamber wall and explore how these intertwined processes in laboratory chambers govern SOA formation and evolution.

A Functional Group Oxidation Model (FGOM) that represents SOA formation and evolution in terms of the competition between functionalization and fragmentation, the extent of oxygen atom addition, and the change of volatility, is developed. The FGOM contains a set of parameters that are to be determined by fitting of the model to laboratory chamber data. The sensitivity of the model prediction to variation of the adjustable parameters allows one to assess the relative importance of various pathways involved in SOA formation.

A critical aspect of the environmental chamber is the presence of the wall, which can induce deposition of SOA-forming vapors and promote heterogeneous reactions. An experimental protocol and model framework are first developed to constrain the vapor-wall interactions. By optimal fitting the model predictions to the observed wall-induced decay profiles of 25 oxidized organic compounds, the dominant parameter governing the extent of wall deposition of a compound is identified, i.e., wall accommodation coefficient. By correlating this parameter with the molecular properties of a compound via its volatility, the wall-induced deposition rate of an organic compound can be predicted based on its carbon and oxygen numbers in the molecule.

Heterogeneous transformation of δ-hydroxycarbonyl, a major first-generation product from long-chain alkane photochemistry, is observed on the surface of particles and walls. The uniqueness of this reaction scheme is the production of substituted dihydrofuran, which is highly reactive towards ozone, OH, and NO3, thereby opening a reaction pathway that is not usually accessible to alkanes. A spectrum of highly-oxygenated products with carboxylic acid, ester, and ether functional groups is produced from the substituted dihydrofuran chemistry, thereby affecting the average oxidation state of the alkane-derived SOA.

The vapor wall loss correction is applied to several chamber-derived SOA systems generated from both anthropogenic and biogenic sources. Experimental and modeling approaches are employed to constrain the partitioning behavior of SOA-forming vapors onto suspended particles vs. chamber walls. It is demonstrated that deposition of SOA-forming vapors to the chamber wall during photooxidation experiments can lead to substantial and systematic underestimation of SOA. Therefore, it is likely that a lack of proper accounting for vapor wall losses that suppress chamber-derived SOA yields contribute substantially to the underprediction of ambient SOA concentrations in atmospheric models.

Habitat segregation in fish assemblages

The segregation of habitats of fish assemblages found in the chalk streams and rivers within the Wessex, South West and Southern Water Authority boundaries in southern England have been examined. Habitat segregation is the most frequent type of resource partitioning in natural communities. The habitat of individual fish species will be defined in order to determine the following: (1) the requirements of each species in terms of depth, current velocity, substrate, cover etc.; (2) identify the essential habitat variables in the segregation of species; (3) whether species in an assemblage demonstrate resource partitioning with reference to habitat, and (4) the mechanisms behind such resource partitioning.

Metallothionein as an indicator of water quality: assessment of the bioavailability of cadmium, copper, mercury and zinc in aquatic animals at the cellular level

The study of metallothioneins (MTs) has greatly improved our understanding of body burdens, metal storage and detoxification in aquatic organisms subjected to contamination by the toxic heavy metals, Cd, Cu, Hg and Zn. These studies have shown that in certain organisms MT status can be used to assess impact of these metals at the cellular level and, whilst validation is currently limited to a few examples, this stress response may be linked to higher levels of organisation, thus indicating its potential for environmental quality assessment. Molluscs, such as Mytilus spp., and several commonly occurring teleost species, are the most promising of the indicator species tested. Natural variability of MT levels caused by the organism's size, condition, age, position in the sexual cycle, temperature and various stressors, can lead to difficulties in interpretation of field data as a definitive response-indicator of metal contamination unless a critical appraisal of these variables is available. From laboratory and field studies these data are almost complete for teleost fish. Whilst for molluscs much of this information is lacking, when suitable controls are utilised and MT measurements are combined with observations of metal partitioning, current studies indicate that they are nevertheless a powerful tool in the interpretation of impact, and may prove useful in water quality assessment.

Mapeamento dos remanescentes de manguezal, a partir da interpretação de ortofotos coloridas, e análise dos vetores de pressão antrópica, como subsídio ao planejamento para conservação do ecossistema no Estado do Rio de Janeiro

Manguezal é um ecossistema costeiro que ocorre nas regiões tropicais e subtropicais do planeta, ocupando a zona entremarés dos oceanos, e sendo caracterizado pela presença de vegetação arbórea adaptada à condições adversas de salinidade, substrato, baixa oxigenação e submersão periódica. A pressão sobre os manguezais do Estado do Rio de Janeiro vem se intensificando nas últimas décadas, e estão associadas a vetores de pressão como os aterros, desmatamentos, queimadas, corte seletivo de madeira, captura predatória de moluscos e crustáceos, lançamento de efluentes de origens diversas, a superexplotação dos recursos pesqueiros e a utilização de técnicas e apetrechos inadequados. Considerando a inexistência de mapeamento integrado e atualizado dos remanescentes de manguezal, indicando sua localização e dimensionamento, o presente estudo veio suprir essa demanda, construindo uma ferramenta consistente para a análise dos principais vetores a que estão expostos, subsidiando a proposição de ações para a conservação e monitoramento desse ecossistema. Essas ações consideram a necessidade de preservação da biodiversidade, da manutenção da atividade pesqueira, da estabilidade da linha de costa, e da subsistência de diversas populações que habitam a região costeira. O mapeamento dos manguezais do Estado do Rio de Janeiro foi elaborado a partir da interpretação visual de ortofotografias coloridas do ano de 2005, na escala 1:10.000, tendo sido realizadas checagens de campo para identificação da verdade terrestre. Os remanescentes mapeados totalizam uma área de aproximadamente 17.720 ha, estando distribuídos por sete regiões hidrográficas localizadas na zona costeira fluminense. Esses ocorrem com mais freqüência, e com maiores dimensões, nas regiões da baía da Ilha Grande, Guandu(Sepetiba) e baía de Guanabara. O estudo contemplou ainda o levantamento e sistematização de dados cartográficos e de sensoriamento remoto, e a identificação e análise dos principais vetores de pressão que atuam sobre esses, a partir da adaptação da metodologia da Análise de Cadeia Causal. Nessa análise foram identificados como principais problemas ambientais dos manguezais fluminenses, a Modificação de habitats e comunidades, a Poluição, e a Exploração não sustentável dos recursos pesqueiros, todos associados aos diferentes vetores de pressão já relacionados. Por fim, foram apresentadas propostas de ações para subsidiar a implementação da Política Estadual para a Conservação dos Manguezais do Estado do Rio de Janeiro, contemplando os níveis operacional, de planejamento, e político. A reativação do Grupo Técnico Permanente sobre Manguezais é de vital importância para a retomada dessas discussões e para a implementação de ações, apoiado na ampliação dos conhecimentos sobre esse rico ecossistema e, integrando e fortalecendo a atuação dos diversos atores envolvidos, buscando assim garantir a integridade dos manguezais fluminenses

Community Turnover of Wood-Inhabiting Fungi across Hierarchical Spatial Scales

For efficient use of conservation resources it is important to determine how species diversity changes across spatial scales. In many poorly known species groups little is known about at which spatial scales the conservation efforts should be focused. Here we examined how the community turnover of wood-inhabiting fungi is realised at three hierarchical levels, and how much of community variation is explained by variation in resource composition and spatial proximity. The hierarchical study design consisted of management type (fixed factor), forest site (random factor, nested within management type) and study plots (randomly placed plots within each study site). To examine how species richness varied across the three hierarchical scales, randomized species accumulation curves and additive partitioning of species richness were applied. To analyse variation in wood-inhabiting species and dead wood composition at each scale, linear and Permanova modelling approaches were used. Wood-inhabiting fungal communities were dominated by rare and infrequent species. The similarity of fungal communities was higher within sites and within management categories than among sites or between the two management categories, and it decreased with increasing distance among the sampling plots and with decreasing similarity of dead wood resources. However, only a small part of community variation could be explained by these factors. The species present in managed forests were in a large extent a subset of those species present in natural forests. Our results suggest that in particular the protection of rare species requires a large total area. As managed forests have only little additional value complementing the diversity of natural forests, the conservation of natural forests is the key to ecologically effective conservation. As the dissimilarity of fungal communities increases with distance, the conserved natural forest sites should be broadly distributed in space, yet the individual conserved areas should be large enough to ensure local persistence.

The juvenile salmonid populations of the River Lune catchment, 1981 to 1985

The aim of this study was to assess the status of the juvenile salmonid populations of the River Lune and its tributaries. There was special emphasis on juvenile salmon stocks in view of the implementation of a net limitation order in 1980. The number of licensed instruments were reduced as follows: (1) Drift, hang or whammel nets - from 12 to 10 (2) Draft or seine nets - from 3 to 1 (3) Heave or haaf nets - from 4 6 to 2 6 For the purpose of this report, the River Lune system has been divided into 13 subcatchments and these are examined with a view to detecting any trends in the data such as subcatchment productivity, partitioning between salmon and trout nursery streams, and whether the restrictions on salmon fishing had any discernible effects on juvenile salmon productivity. The effect of flow and instream obstructions on salmonid densities are also investigated as these have been shown to be important factors affecting abundance. Throughout the study period a programme of enhancement stocking took place primarily with salmon ova and fry. The possible impact of this on the results of the surveys has been assessed. The distribution and abundance of juvenile salmon was found to be relatively consistent in each of the 13 subcatchments studied over the 1981-1985 period.

Postrelease survival, vertical and horizontal movements, and thermal habitats of five species of pelagic sharks in the central Pacific Ocean

From 2001 to 2006, 71 pop-up satellite archival tags (PSATs) were deployed on five species of pelagic shark (blue shark [Prionace glauca]; shortfin mako [Isurus oxyrinchus]; silky shark [Carcharhinus falciformis]; oceanic whitetip shark [C. longimanus]; and bigeye thresher [Alopias superciliosus]) in the central Pacific Ocean to determine species-specific movement patterns and survival rates after release from longline fishing gear. Only a single postrelease mortality could be unequivocally documented: a male blue shark which succumbed seven days after release. Meta-analysis of published reports and the current study (n=78 reporting PSATs) indicated that the summary effect of postrelease mortality for blue sharks was 15% (95% CI, 8.5–25.1%) and suggested that catch-and-release in longline fisheries can be a viable management tool to protect parental biomass in shark populations. Pelagic sharks displayed species-specific depth and temperature ranges, although with significant individual temporal and spatial variability in vertical movement patterns, which were also punctuated by stochastic events (e.g., El Niño-Southern Oscillation). Pelagic species can be separated into three broad groups based on daytime temperature preferences by using the unweighted pair-group method with arithmetic averaging clustering on a Kolmogorov-Smirnov Dmax distance matrix: 1) epipelagic species (silky and oceanic whitetip sharks), which spent >95% of their time at temperatures within 2°C of sea surface temperature; 2) mesopelagic-I species (blue sharks and shortfin makos, which spent 95% of their time at temperatures from 9.7° to 26.9°C and from 9.4° to 25.0°C, respectively; and 3) mesopelagic-II species (bigeye threshers), which spent 95% of their time at temperatures from 6.7° to 21.2°C. Distinct thermal niche partitioning based on body size and latitude was also evident within epipelagic species.

Distribuição das assembleias de girinos associadas a riachos da Reserva Ecológica de Guapiaçu, Cachoeiras de Macacú, Rio de Janeiro, Brasil

Os girinos são organismos diversos e abundantes nos pequenos riachos de cabeceira de florestas tropicais e constituem importantes componentes da diversidade biológica, da trófica e funcional dos sistemas aquáticos. Diferentes características estruturais e limnológicas dos ambientes aquáticos influenciam a organização das assembleias de girinos. Embora o estágio larvar dos anuros seja o mais vulnerável de seu ciclo de vida, sujeito a elevadas taxas de mortalidade, as pesquisas sobre girinos na região neotropical ainda são pouco representativas diante da elevada diversidade de anfíbios desta região e ferramentas que permitam a sua identificação ainda são escassas. Nesta tese, dividida em três capítulos, apresento uma compilação das informações relacionadas aos principais fatores que afetam as assembleias de girinos na região tropical (Capítulo 1), a caracterização morfológica dos girinos encontrados nos riachos durante o estudo e uma proposta de chave dicotômica de identificação (Capítulo 2) e avalio a importância relativa da posição geográfica e da variação temporal de fatores ambientais locais sobre as assembleias de girinos, assim como a correlação entre as espécies de girinos e as variáveis ambientais de 10 riachos, ao longo de 15 meses, nas florestas da REGUA (Capítulo 3). Há pelo menos oito tendências relacionadas à distribuição das assembleias de girinos: (1) o tamanho dos riachos e a diversidade de microhabitats são importantes características abióticas influenciando a riqueza e a composição de espécies; (2) em poças, o gradiente de permanência (e.g., hidroperíodo) e a heterogeneidade do habitat são os principais fatores moldando as assembleias de girinos; (3) a composição de espécies parece ser um parâmetro das assembleias mais relevante do que a riqueza de espécies e deve ser primeiramente considerado durante o planejamento de ações conservacionistas de anuros associados a poças e riachos; (4) a predação parece ser a interação biótica mais importante na estruturação das assembleias de girinos, com predadores vertebrados (e.g. peixes) sendo mais vorazes em habitats permanentes e predadores invertebrados (e.g. larvas de odonata) sendo mais vorazes em ambientes temporários; (5) os girinos podem exercer um efeito regulatório, predando ovos e girinos recém eclodidos; (6) o uso do microhabitat varia em função da escolha do habitat reprodutivo pelos adultos, presença de predadores, filogenia, estágio de desenvolvimento e heterogeneidade do habitat; (7) os fatores históricos restringem os habitats reprodutivos que uma espécie utiliza, impondo restrições comportamentais e fisiológicas; (8) a variação temporal nos fatores bióticos (e.g., fatores de risco), abióticos (e.g., distribuição de chuvas), e no padrão de reprodução das espécies pode interferir na estrutura das assembleias de girinos tropicais. A variação temporal na heterogeneidade ambiental dos riachos da REGUA resultou na previsibilidade das assembleias locais de girinos, sendo que os parâmetros ambientais explicaram 23% da variação na sua composição. Os parâmetros espaciais explicaram uma porção menor da variação nas assembleias (16%), enquanto uma porção relativamente elevada da variação temporal da heterogeneidade ambiental foi espacialmente estruturada (18%). As variáveis abióticas que apresentaram as maiores correlação com a composição das assembleias de girinos foram a proporção de folhiço e de rochas no fundo do riacho, e secundariamente a profundidade, a condutividade e a temperatura. O gradiente gerado pela proporção de folhiço e de rochas representou a transição entre riachos permanentes e intermitentes. Este gradiente proporcionou o turnover de espécies, o qual também seguiu um gradiente de condutividade, temperatura, profundidade, e em menor extensão, de hidroperíodo e largura, que estiveram fortemente associado ao grau de permanência dos riachos. Estes resultados corroboram tanto a hipótese do controle ambiental, como do controle biótico de comunidades e indicam que a variação temporal da heterogeneidade ambiental e a variação na posição geográfica são importantes para a estruturação local de assembleias de girinos da REGUA. Os resultados também permitiram distinguir entre assembleias de girinos exclusivas de riachos permanentes, exclusivas de riachos intermitentes e aquelas registradas nos dois tipos de riachos. Os resultados deste capítulo são relevantes para compreender em que extensão os efeitos da variação temporal na heterogeneidade ambiental e de processos espaciais afetam localmente a estruturação de assembleias de girinos.

Testes de sistemas de informações geográficas com lógica nebulosa.

Os testes são uma atividade crucial no desenvolvimento de sistemas, pois uma boa execução dos testes podem expor anomalias do software e estas podem ser corrigidas ainda no processo de desenvolvimento, reduzindo custos. Esta dissertação apresenta uma ferramenta de testes chamada SIT (Sistema de Testes) que auxiliará no teste de Sistemas de Informações Geográficas (SIG). Os SIG são caracterizados pelo uso de informações espaciais georreferenciadas, que podem gerar um grande número de casos de teste complexos. As técnicas tradicionais de teste são divididas em funcionais e estruturais. Neste trabalho, o SIT abordará os testes funcionais, focado em algumas técnicas clássicas como o particionamento de equivalência e análise do Valor Limite. O SIT também propõe o uso de Lógica Nebulosa como uma ferramenta que irá sugerir um conjunto mínimo de testes a executar nos SIG, ilustrando os benefícios da ferramenta.

Multiview stereo via volumetric Graph-Cuts and occlusion robust photo-consistency.

This paper presents a volumetric formulation for the multi-view stereo problem which is amenable to a computationally tractable global optimisation using Graph-cuts. Our approach is to seek the optimal partitioning of 3D space into two regions labelled as "object" and "empty" under a cost functional consisting of the following two terms: (1) A term that forces the boundary between the two regions to pass through photo-consistent locations and (2) a ballooning term that inflates the "object" region. To take account of the effect of occlusion on the first term we use an occlusion robust photo-consistency metric based on Normalised Cross Correlation, which does not assume any geometric knowledge about the reconstructed object. The globally optimal 3D partitioning can be obtained as the minimum cut solution of a weighted graph.

Distribution and co-occurrence of rockfishes (family: Sebastidae) over trawlable shelf and slope habitats of California and southern Oregon

The rockfishes of the sebastid genus Sebastes are a very important fishery resource off the coasts of California and southern Oregon. How-ever, many of the 54 managed stocks of west coast rockfish have recently reached historically low population levels, leading fishery managers to re-examine current management practices. Management of rockfish stocks as multispecies aggregates, as opposed to independent stocks within the ground-fish fishery, can be more desirable when nontargeted bycatch, discard, and management complexity are considered. Rockfish assemblage structure and species co-occurrences were determined by using data from the Alaska Fisheries Science Center triennial continental shelf bottom trawl survey. The weight of rockfish species in trawl catches was expressed as a catch-per-unit-of-effort (CPUE) statistic, from which species spatial distributions, overlaps, diversity, and richness were analyzed. Multidimensional scaling of transformed CPUE data was employed in indirect gradient and multivariate partitioning analyses to quantify assemblage relationships. Results indicated that rockfish distributions closely match the bathymetry of coastal waters. Indirect gradient analysis suggested that depth and latitude are the principal factors in structuring the spatial distributions of rockfish on trawlable habitat. In addition, four assemblages were identified through the joint evaluation of species’ distributions and multivariate partitioning analyses: 1) deep-water slope; 2) northern shelf; 3) southern shelf; and 4) nearshore. The slope, shelf, and near-shore groups are found in depth ranges of 200–500 m, 100–250 m, and 50–150 m, respectively. The division of northern and southern shelf assemblages occurs over a broad area between Cape Mendocino and Monterey Canyon. The results of this analysis are likely to have direct application in the management of rockfish stocks off the coasts of southern Oregon and California.

古田山常绿阔叶林物种生境关联及其对物种共存的贡献

物种共存机制一直是群落生态学研究的核心内容。解释物种共存的假说很多，近年来最引人注目的是生态位分化假说和群落中性理论。这两种理论对群落内物种共存的相对重要性是目前群落生态学研究的热点。国际上这方面的研究基本集中在热带森林大样地内，而针对亚热带森林大样地的相关研究却非常少见。本文以浙江古田山24公顷常绿阔叶林永久固定监测样地第一次调查数据为基础，研究样地内木本植物与生境关联和与地形因子梯度相关，目的在于探讨物种生境生态位分化在亚热带常绿阔叶林内物种多样性维持中的作用，也为了解决当前物种生境关联相关研究中的一些不足之处。 本研究首先采用Torus转换检验分析古田山样地内90种常见木本植物与5类生境关联，结果表明有75种（83.3%）至少与一类生境类型显著相关，说明大部分亚热带森林群落内物种具有生境特化的特性。与CTFS全球大样地类似研究结果相比，古田山样地内生境特化的物种比例更高，这与古田山样地复杂的地形条件密切相关。通过本研究，证明地形条件越复杂，物种特化比例越高的趋势确实存在。在古田山样地，虽然83.3%的物种有生境特化的特性，但生境特化没有排他性，即并不完全排斥其他生境，在非最适生境也能稳定与其他物种共存。根据本研究与生境负相关的平均物种数与被检测的物种比例估算，古田山样地内生境异质性对物种多样性维持的贡献率约为19.6%，说明物种生境特化对于物种共存有一定的作用，但贡献并不大，除了生境异质性，仍有其他因素决定物种共存。 目前有关物种生境关联的研究都假设同一物种的个体对于生境偏好一致，不管胸径大小是否相同，很少有人研究不同生长阶段生境偏好的变化。本研究利用Torus转换检验比较样地内60种常见木本植物在3个生活史阶段（幼苗阶段、小树阶段和成熟阶段）与5类生境关联的变化，结果表明大部分物种在其幼苗和小树阶段的生境偏好比较一致，但成熟阶段与前两个阶段差别比较大，说明物种在生活史不同阶段的生境偏好可能发生改变。 由于生境划分没有统一的标准，不同学者用不同的方法划分生境类型和数量，主观性很大，造成不同研究地点、不同研究者之间的研究结果可比性差。本研究尝试利用直接梯度分析方法重新分析物种空间分布与地形梯度相关，用以代替生境关联分析。利用CCA分析检验样地内90种常见木本植物与4种地形因子梯度（海拔、凹凸度、坡度和坡向）的相关情况，发现有76个（84%）物种的空间分布与地形梯度相关，说明大部分物种具有沿地形因子梯度分布的特性。4种地形因子梯度变化对90个物种空间分布的解释量约为20%，说明地形异质性对于物种共存有一定作用，但贡献并不大。对生境关联分析和直接梯度法分析结果进行比较，两种方法分析结果一致，得到的结论也一致。直接梯度分析方法可以避免生境划分对研究结果的影响，是今后类似研究中值得采纳的方法。 本研究的结果表明地形异质性引起的生态位分化在古田山样地群落内物种多样性维持中起一定的作用，但贡献不大，因此需要更深入地研究其他环境因素引起的生态位分化在物种多样性维持中的作用，同时也需要进一步研究中性过程在物种多样性维持中的作用，以更全面地探讨生态位分化假说和中性理论在亚热带常绿阔叶林内物种共存的相对重要性。

Novel architecture to facilitate disassembly and reuse of electronic components and sub-assemblies

This paper advances the proposition that in many electronic products, the partitioning scheme adopted and the interconnection system used to interconnect the sub-assemblies or components are intimately related to the economic benefits, and hence the attractiveness, of reuse of these items. An architecture has been developed in which the residual values of the connectors, components and sub-assemblies are maximized, and opportunities for take-back and reuse of redundant items are greatly enhanced. The system described also offers significant manufacturing cost benefits in terms of ease of assembly, compactness and robustness.

Online multiple classifier boosting for object tracking

This paper presents a new online multi-classifier boosting algorithm for learning object appearance models. In many cases the appearance model is multi-modal, which we capture by training and updating multiple strong classifiers. The proposed algorithm jointly learns the classifiers and a soft partitioning of the input space, defining an area of expertise for each classifier. We show how this formulation improves the specificity of the strong classifiers, allowing simultaneous location and pose estimation in a tracking task. The proposed online scheme iteratively adapts the classifiers during tracking. Experiments show that the algorithm successfully learns multi-modal appearance models during a short initial training phase, subsequently updating them for tracking an object under rapid appearance changes. © 2010 IEEE.

Diffusive coevolution and mutualism maintenance mechanisms in a fig-fig wasp system

In reciprocal mutualism systems, the exploitation events by exploiters might disrupt the reciprocal mutualism, wherein one exploiter species might even exclude other coexisting exploiter species over an evolutionary time frame. What remains unclear is how such a community is maintained. Niche partitioning, or spatial heterogeneity among the mutualists and exploiters, is generally believed to enable stability within a mutualistic system. However, our examination of a reciprocal mutualism between a fig species (Ficus racemosa) and its pollinator wasp (Ceratosolen fusciceps) shows that spatial niche partitioning does not sufficiently prevent exploiters from overexploiting the common resource (i.e., the female flowers), because of the considerable niche overlap between the mutualists and exploiters. In response to an exploiter, our experiment shows that the fig can (1) abort syconia-containing flowers that have been galled by the exploiter, Apocryptophagus testacea, which oviposits before the pollinators do; and (2) retain syconia-containing flowers galled by Apocryptophagus mayri, which oviposit later than pollinators. However, as a result of (2), there is decreased development of adult non-pollinators or pollinator species in syconia that have not been sufficiently pollinated, but not aborted. Such discriminative abortion of figs or reduction in offspring development of exploiters while rewarding cooperative individuals with higher offspring development by the fig will increase the fitness of cooperative pollinating wasps, but decrease the fitness of exploiters. The fig fig wasp interactions are diffusively coevolved, a case in which fig wasps diversify their genotype, phenotype, or behavior as a result of competition between wasps, while figs diverge their strategies to facilitate the evolution of cooperative fig waps or lessen the detrimental behavior by associated fig wasps. In habitats or syconia that suffer overexploitation, discriminative abortion of figs or reduction in the offspring development of exploiters in syconia that are not or not sufficiently pollinated will decrease exploiter fitness and perhaps even drive the population of exploiters to local extinction, enabling the evolution and maintenance of cooperative pollinators through the movement between habitats or syconia (i.e., the metapopulations).