In natural systems, it has been observed that plankton exist in patches rather than in an even distribution across a body of water. However, the mechanisms behind this patchiness are not fully understood. Several previous modeling studies have examined the effects of abiotic and biotic factors on patch structure. Yet these models ignore a key point: zooplankton often undergo diel vertical migration. I have formulated a model that incorporates vertical movement into the Rosezweig-MacArthur (R-M) predator-prey model. The R-M model is stable only at a carrying capacity below a critical value. I found that adding vertical movement stabilizes the system even at a high carrying capacity. By analyzing temporal stability and spatial structure, my results show that vertical movement interacts with carrying capacity to determine patch structure. / by Simantha Ather. / Thesis (M.S.)--Florida Atlantic University, 2009. / Includes bibliography. / Electronic reproduction. Boca Raton, Fla., 2009. Mode of access: World Wide Web.

O virioplâncton é o mais diverso e o mais abundante componente do plâncton, o qual exerce grande importância ecológica, estando envolvido em processos de ciclagem de nutrientes, diversificação e distribuição de algas e bactérias, controle de florações fitoplanctônicas, formação de nuvens e transferência gênica horizontal. Os objetivos deste projeto foram analisar as populações das famílias Phycodnaviridae e Myoviridae, nos rios Cuieiras, Negro e Solimões (AM), e determinar um possível impacto do lançamento de efluente doméstico sobre essas populações. Miofagos foram detectados nas amostras dos 3 rios estudados, enquanto que os ficodnavírus somente foram encontrados nos rios Solimões e Cuieiras. Análises filogenéticas revelaram que os miofagos não se agrupam em função das características do ambiente nem da localidade, diferentemente dos ficodnavírus. A interferência do lançamento de efluentes domésticos sobre as características da água e sobre as populações locais das famílias Phycodnaviridae e Myoviridae e fitoplanctônicas não foi observada. / The virioplankton is the most diverse and abundant plankton component, which exerts great ecological importance by being involved in processes such as nutrient cycling, diversification and distribution of algae and bacteria, bloom termination, cloud formation and horizontal gene transfer. The aims of the present work were to analyze the viral populations of Phycodnaviridae and Myoviridae families from the Cuieiras, Negro and Solimões rivers (AM), and to determine whether or not sewage discharge in the area is affecting these populations. Myophages were detected in all samples from the 3 studied rivers while phycodnaviruses were only found in the Solimões and Cuieiras rivers. Phylogenetic analyses revealed that myophages do not cluster accordingly neither to the environment nor to the location, differently from phycodnaviruses. The interference of sewage discharge on the water characteristics and the local viral (phycodnavirus and myophage) and phytoplanktonic populations was not detected.

Neste trabalho foi desenvolvida uma instrumentação óptica in-line para filmagem de partículas in situ, utilizando a técnica de sombreamento, acompanhada de softwares para processamento das imagens. As novas ferramentas foram implementadas e foi conduzido um estudo de caso para a avaliação da dinâmica de partículas e suas forçantes ambientais em uma região costeira de Ubatuba. O sistema com 4 sensores acoplados, adquiriu dados oceanográficos e de imagens a cada 40 minutos, no período entre 20 de fevereiro e 7 de junho de 2017. Dados metereológicos adicionais foram obtidos para análises correlativas. Testes com os softwares de visão computacional estabeleceram o uso dos filtros de tamanho entre 500 e 16300 pixels e contraste maior que 50% para segmentação das Regiões de Interesse (ROIs). Os testes também indicaram a melhor performance do algoritmo de segmentação ModeValue e de uma base de treinamento composta por 9 classes com 300 organismos cada para a classificação automática. Devido à baixa acurácia obtida na etapa de classificação automática de imagens desconhecidas de organismos (27%), os objetos foram analisados nesse estudo como partículas, divididos de acordo com o tamanho de seu eixo maior em três ranges (<385 μm; 385-620 μm; >620μm). A análise das partículas, por questões estatísticas, considerou o maior intervalo de amostragens contínuas da série, de 5 de abril a 7 de maio de 2017. Primeiramente a série de cada variável foi decomposta em componentes harmônicas, com base na análise de Fourier, visando detectar padrões recorrentes e, em seguida, as variáveis com picos de densidade espectral mais expressivos em frequências da ordem de uma semana e de um dia foram comparadas por meio da coerência quadrada. As partículas menores, do range 1, foram as que melhor representaram a abundância total e os maiores valores de coerência foram entre as variáveis com período característico de um dia. A instrumentação implementada e testada ao longo de cerca de 3 meses possibilitou a aquisição de imagens de qualidade e de resultados iniciais úteis para o aprimoramento do sistema, visando instalações futuras de longo prazo em ambientes costeiros. / In this work, in-line optical optical instrumentation was developed for in situ particle filming, using the shadowgraphic technique, accompanied by image processing software. The new tools were implemented and a case study was carried out to evaluate the dynamics of particles and their environmental forcing in a coastal region of Ubatuba. The system, with 4 coupled environmental sensors, acquired oceanographic and image data every 40 minutes, between February 20 and June 7, 2017. Aditional meteorological data were obtained for correlative analyzes. Tests with the implemented computer vision software have stablished the use of size filters between 500 and 16300 pixels and contrast level greater than 50% for the segmentation of Regions of Interest (ROIs). The test also indicated the best performance of the Mode Value segmentation algorithm and of a training set composed by 9 classes with 300 organisms each for automatic classification. Due to the low global accuracy obtained in the automatic classification stage of unknown images of organisms (27%), the objects were analyzed in this study as particle, divided according to the size of their major axis in three ranges (<385 μm; 385 - 620 μm; >620μm). Particle analysis, for statistical reasons, considered the largest continuous sampling range of the series, from April 5 to May 7, 2017. First, the series of each variables decomposed into harmonic components, based on Fourier analysis, aiming to detect recurrent patterns, and then the variables with more expressive spectral density peaks at frequencies of the order of one week and one day were compared by means of square coherence. The smaller particles of range 1 were the ones that best represented the total abundance, and the highest values of coherence were among the variables with characteristic period of one day. The instrumentation implemented and tested over about 3 months allowed the acquisition og high-quality images and the initial results were useful for improving the system, aiming at future long-term deployments in coastal environments.

This thesis is centered on the application of the Normalized Biomass Size Spectra (NBSS) theory to the study of plankton systems in shelf and oceanic areas of the tropical and subtropical Southwest Atlantic. I evaluated NBSS parameters over different environmental settings and their utility as proxies for system in Brazilian waters. The LOPC and the ZooScan are recently developed optical systems to automatically detect and measure plankton and particle size distributions in situ and in laboratory, respectively. I present two case studies: the first deals with the spatial variability over the Abrolhos Bank and adjacent oceanic areas in tropical latitudes, and the second focuses on temporal variability of plankton communities along a 5-year time series on a fixed station on the inner shelf at a subtropical location (Ubatuba, São Paulo). The data sets consisted of vertical profiles obtained with a LOPC and plankton preserved samples collected with a 200-m-mesh net. I observed in both data sets the accumulation of small particles (< 1 mm) above and within the pycnocline. NBSS slopes and intercepts were significantly different according to the contrasting environmental conditions observed in both areas; higher intercepts and steeper slopes were associated with higher productivity. The results highlight: i) water-column stratification as a key feature driving particle and plankton vertical distribution, ii) NBSS parameters as indicators of different environmental settings, and iii) that the cold and nutrient-rich South Atlantic Central Water (SACW) have an important role structuring the zooplankton size distributions over the Abrolhos Bank and vicinities and off Ubatuba. The NBSS parameters associated with information on plankton composition and distributions provided important information to evaluate the influence of oceanographic forcing on plankton dynamics in distinct ecosystems of the Southwest Atlantic / O tema central desta tese é a aplicação da teoria do espectro de biomassa normalizado (NBSS, em inglês) no estudo de sistemas planctônicos em áreas costeiras e oceânicas, no Atlântico Sudoeste tropical e subtropical. Eu avaliei os parâmetros do NBSS em diferentes situações ambientais na Plataforma Continental Brasileira e a utilidade desses índices para inferir sobre produtividade. O LOPC e o ZooScan são sistemas ópticos recentemente desenvolvidos para detectar e mensurar a distribuição de tamanhos de partículas e zooplâncton in situ e em laboratório, respectivamente. Eu apresento dois estudos de caso: o primeiro lida com a variabilidade espacial sobre o banco de Abrolhos e áreas adjacentes oceânicas em latitudes tropicais, enquanto o segundo enfatiza a variabilidade temporal de comunidades planctônicas ao longo de 5 anos numa estação fixa na plataforma interna em uma localidade subtropical (Ubatuba, SP). O conjunto de dados consistiu de perfis verticais obtidos com o LOPC e com rede de plâncton, malha de 200-m. Eu observei o acúmulo de partículas < 1 mm na picnoclina e acima desta de forma consistente. Ambos inclinação e intercepto das retas ajustadas ao NBSS responderam à condições ambientais contrastantes em ambas as áreas; associamos interceptos maiores e inclinações mais negativas com maior produtividade do plâncton. Os principais resultados indicam: i) que a estratificação é um fator chave na distribuição vertical de partículas e do plâncton, ii) os parâmetros do NBSS como indicadores de diferentes condições ambientais e iii) que a Água Central do Atlântico Sul (ACAS, fria e rica em nutrientes) tem um importante papel na estruturação da distribuição de tamanhos acima do Banco de Abrolhos e proximidades e ao largo de Ubatuba. Os parâmetros do NBSS, associados com informações sobre a composição taxonômica e distribuição do plâncton foram importantes para avaliar a influência de feições oceanográficas sobre a dinâmica do mesozooplâncton em diferentes ecossistemas no Atlântico Sudoeste

Neste trabalho foi desenvolvida uma instrumentação óptica in-line para filmagem de partículas in situ, utilizando a técnica de sombreamento, acompanhada de softwares para processamento das imagens. As novas ferramentas foram implementadas e foi conduzido um estudo de caso para a avaliação da dinâmica de partículas e suas forçantes ambientais em uma região costeira de Ubatuba. O sistema com 4 sensores acoplados, adquiriu dados oceanográficos e de imagens a cada 40 minutos, no período entre 20 de fevereiro e 7 de junho de 2017. Dados metereológicos adicionais foram obtidos para análises correlativas. Testes com os softwares de visão computacional estabeleceram o uso dos filtros de tamanho entre 500 e 16300 pixels e contraste maior que 50% para segmentação das Regiões de Interesse (ROIs). Os testes também indicaram a melhor performance do algoritmo de segmentação ModeValue e de uma base de treinamento composta por 9 classes com 300 organismos cada para a classificação automática. Devido à baixa acurácia obtida na etapa de classificação automática de imagens desconhecidas de organismos (27%), os objetos foram analisados nesse estudo como partículas, divididos de acordo com o tamanho de seu eixo maior em três ranges (&#60;385 &#956;m; 385-620 &#956;m; &#62;620&#956;m). A análise das partículas, por questões estatísticas, considerou o maior intervalo de amostragens contínuas da série, de 5 de abril a 7 de maio de 2017. Primeiramente a série de cada variável foi decomposta em componentes harmônicas, com base na análise de Fourier, visando detectar padrões recorrentes e, em seguida, as variáveis com picos de densidade espectral mais expressivos em frequências da ordem de uma semana e de um dia foram comparadas por meio da coerência quadrada. As partículas menores, do range 1, foram as que melhor representaram a abundância total e os maiores valores de coerência foram entre as variáveis com período característico de um dia. A instrumentação implementada e testada ao longo de cerca de 3 meses possibilitou a aquisição de imagens de qualidade e de resultados iniciais úteis para o aprimoramento do sistema, visando instalações futuras de longo prazo em ambientes costeiros. / In this work, in-line optical optical instrumentation was developed for in situ particle filming, using the shadowgraphic technique, accompanied by image processing software. The new tools were implemented and a case study was carried out to evaluate the dynamics of particles and their environmental forcing in a coastal region of Ubatuba. The system, with 4 coupled environmental sensors, acquired oceanographic and image data every 40 minutes, between February 20 and June 7, 2017. Aditional meteorological data were obtained for correlative analyzes. Tests with the implemented computer vision software have stablished the use of size filters between 500 and 16300 pixels and contrast level greater than 50% for the segmentation of Regions of Interest (ROIs). The test also indicated the best performance of the Mode Value segmentation algorithm and of a training set composed by 9 classes with 300 organisms each for automatic classification. Due to the low global accuracy obtained in the automatic classification stage of unknown images of organisms (27%), the objects were analyzed in this study as particle, divided according to the size of their major axis in three ranges (&#60;385 &#956;m; 385 - 620 &#956;m; &#62;620&#956;m). Particle analysis, for statistical reasons, considered the largest continuous sampling range of the series, from April 5 to May 7, 2017. First, the series of each variables decomposed into harmonic components, based on Fourier analysis, aiming to detect recurrent patterns, and then the variables with more expressive spectral density peaks at frequencies of the order of one week and one day were compared by means of square coherence. The smaller particles of range 1 were the ones that best represented the total abundance, and the highest values of coherence were among the variables with characteristic period of one day. The instrumentation implemented and tested over about 3 months allowed the acquisition og high-quality images and the initial results were useful for improving the system, aiming at future long-term deployments in coastal environments.

This thesis is centered on the application of the Normalized Biomass Size Spectra (NBSS) theory to the study of plankton systems in shelf and oceanic areas of the tropical and subtropical Southwest Atlantic. I evaluated NBSS parameters over different environmental settings and their utility as proxies for system in Brazilian waters. The LOPC and the ZooScan are recently developed optical systems to automatically detect and measure plankton and particle size distributions in situ and in laboratory, respectively. I present two case studies: the first deals with the spatial variability over the Abrolhos Bank and adjacent oceanic areas in tropical latitudes, and the second focuses on temporal variability of plankton communities along a 5-year time series on a fixed station on the inner shelf at a subtropical location (Ubatuba, São Paulo). The data sets consisted of vertical profiles obtained with a LOPC and plankton preserved samples collected with a 200-m-mesh net. I observed in both data sets the accumulation of small particles (< 1 mm) above and within the pycnocline. NBSS slopes and intercepts were significantly different according to the contrasting environmental conditions observed in both areas; higher intercepts and steeper slopes were associated with higher productivity. The results highlight: i) water-column stratification as a key feature driving particle and plankton vertical distribution, ii) NBSS parameters as indicators of different environmental settings, and iii) that the cold and nutrient-rich South Atlantic Central Water (SACW) have an important role structuring the zooplankton size distributions over the Abrolhos Bank and vicinities and off Ubatuba. The NBSS parameters associated with information on plankton composition and distributions provided important information to evaluate the influence of oceanographic forcing on plankton dynamics in distinct ecosystems of the Southwest Atlantic / O tema central desta tese é a aplicação da teoria do espectro de biomassa normalizado (NBSS, em inglês) no estudo de sistemas planctônicos em áreas costeiras e oceânicas, no Atlântico Sudoeste tropical e subtropical. Eu avaliei os parâmetros do NBSS em diferentes situações ambientais na Plataforma Continental Brasileira e a utilidade desses índices para inferir sobre produtividade. O LOPC e o ZooScan são sistemas ópticos recentemente desenvolvidos para detectar e mensurar a distribuição de tamanhos de partículas e zooplâncton in situ e em laboratório, respectivamente. Eu apresento dois estudos de caso: o primeiro lida com a variabilidade espacial sobre o banco de Abrolhos e áreas adjacentes oceânicas em latitudes tropicais, enquanto o segundo enfatiza a variabilidade temporal de comunidades planctônicas ao longo de 5 anos numa estação fixa na plataforma interna em uma localidade subtropical (Ubatuba, SP). O conjunto de dados consistiu de perfis verticais obtidos com o LOPC e com rede de plâncton, malha de 200-m. Eu observei o acúmulo de partículas < 1 mm na picnoclina e acima desta de forma consistente. Ambos inclinação e intercepto das retas ajustadas ao NBSS responderam à condições ambientais contrastantes em ambas as áreas; associamos interceptos maiores e inclinações mais negativas com maior produtividade do plâncton. Os principais resultados indicam: i) que a estratificação é um fator chave na distribuição vertical de partículas e do plâncton, ii) os parâmetros do NBSS como indicadores de diferentes condições ambientais e iii) que a Água Central do Atlântico Sul (ACAS, fria e rica em nutrientes) tem um importante papel na estruturação da distribuição de tamanhos acima do Banco de Abrolhos e proximidades e ao largo de Ubatuba. Os parâmetros do NBSS, associados com informações sobre a composição taxonômica e distribuição do plâncton foram importantes para avaliar a influência de feições oceanográficas sobre a dinâmica do mesozooplâncton em diferentes ecossistemas no Atlântico Sudoeste

Thesis: Ph. D., Joint Program in Biological Oceanography (Massachusetts Institute of Technology, Department of Biology; and the Woods Hole Oceanographic Institution), 2017. / Cataloged from PDF version of thesis. / Includes bibliographical references (pages 159-168). / Thecosome pteropods are planktonic mollusks that form aragonite shells and that may experience increased dissolution and other adverse effects due to ocean acidification. This thesis focuses on assessing the possible biological effects of ocean acidification on the shells and locomotion of pteropods and examining the response of a local pteropod population to environmental change over time. I analyzed shell condition after exposing pteropods to elevated CO₂ as well as in natural populations to investigate the sensitivity of the shells of different species to aragonite saturation state ([omega][subscript A]). The pteropods (Limacina retroversa) from laboratory experiments showed the clearest pattern of shell dissolution in response to decreased [omega][subscript A], while wild populations either had non-significant regional trends in shell condition (Clio pyramidata) or variability in shell condition that did not match expectations due to regional variability in [omega][subscript A] (Limacina helicina). At locations with intermediate [omega][subscript A] (1.5-2.5) the variability seen in L. helicina shell condition might be affected by food availability more than tA. I examined sinking and swimming behaviors in the laboratory in order to investigate a possible fitness effect of ocean acidification on pteropods. The sinking rates of L. retroversa from elevated CO₂ treatments were slower in conjunction with worsened shell condition. These changes could increase their vulnerability to predators in the wild. Swimming ability was mostly unchanged by elevated CO₂ after experiments that were up to three weeks in duration. I used a long-term dataset of pteropods in the Gulf of Maine to directly test whether there has been a population effect of environmental change over the past several decades. I did not observe a population decline between 1977 and 2015, and L. retroversa abundance in the fall actually increased over the time series. Analysis of the habitat use of L. retroversa revealed seasonal associations with temperature, salinity, and bottom depths. The combination of laboratory experiments and field surveys helped to address gaps in knowledge about pteropod ecology and improve our understanding of the effects of ocean acidification on pteropods. / by Alexander Bergan. / Ph. D.

In the summer of 1932, an investigation of some biological, physical, and chemical conditions at the Bear River Migratory Waterfowl Refuge, Utah, was begun at the Utah State Agricultural College to determine some of the factors governing the biological productivity of brackish-water marshes. Unfortunately, lack of funds made it impossible to continue work beyond the first season.

The assemblage composition, biomass and dynamics of zooplankton and epibenthos were examined in a commercial prawn pond in southeast Queensland over two seasons. Physico-chemical characteristics of the pond water were measured concurrently. Numbers and biomass of zooplankton in the surface tows (140 micrometre mesh) varied from 8 ind. L-1 (44 micrograms L-1) to 112 ind. L-1 (324 micrograms L-1) in the first season, with peaks in biomass corresponding to peaks in numbers. In the second season the zooplankton numbers varied from 12 to 590 ind. L-1, but peaks in numbers did not correspond with peaks in biomass, which varied from 28 to 465 micrograms L-1. This was due to differences in the size of the dominant taxa across the season. Although this occurred in both seasons, the effect on biomass was more pronounced in the second season. In both seasons, immediately after the ponds were stocked with prawn postlarvae there was a rapid decline in zooplankton numbers, particularly of the dominant larger copepods. This was probably due to predation by the postlarvae. Subsequent peaks in zooplankton numbers were principally due to barnacle nauplii. The largest peaks in zooplankton numbers occurred before stocking in the first season, but the largest peaks were in the middle of the second season. While changes in abundance and biomass of the zooplankton assemblage were not correlated with physico-chemical characteristics in the first season, there were correlations between zooplankton numbers and temperature, dissolved oxygen, pH and secchi disk readings in the second season. No correlations were found with zooplankton biomass and physico-chemical characteristics in the second season. The correlations in the second season were mainly due to the high prevalence of barnacle nauplii through the middle part of the season, and may reflect suitable conditions for barnacle reproduction. Epibenthic faunal abundance in the beam trawls (1 mm mesh) peaked at 14 ind. m-2 and 7 ind. m-2 in the first and second seasons respectively and the biomasses at 0.8 g m-2 and 0.7 g m-2. Peaks in abundance of epibenthos did not correspond to peaks in biomass. This was due to large differences in the size of the taxa across the seasons. Sergestids (Acetes sibogae) and amphipods were the most abundant taxa in beam trawl samples. Amphipods were only abundant in the first season, with their numbers increasing towards the end of the grow out period. Acetes were abundant in both seasons, but were dominant in the second season. Correlations between physico-chemical parameters and epibenthos numbers were found to be strongly influenced by the dominant taxa in each season. In the first season, negative correlations were found between epibenthos abundance and pH and temperature. These relationships may reflect an effect on the growth of macroalgae in the pond, with which the amphipods were strongly associated, rather than a direct effect on the epibenthos. In the second season, a positive correlation existed between temperature and epibenthos abundance, however this was strongly influenced by the very high abundance of Acetes in the last sampling period. No correlations were found between epibenthic fauna biomass and physico-chemical parameters. Abundances of epibenthic fauna were not related to zooplankton densities indicating this source of food was not likely to be a limiting factor. Neither the pond water exchange regime nor moon phase could explain changes observed in abundances of zooplankton or epibenthos assemblages in the first season, however the sampling regime was not designed to specifically investigate these effects. In the second season water exchanges were sampled more rigorously. The density of zooplankton in the outlet water was from 2 to 59% of the density of zooplankton in the pond, and the zooplankton density of the inlet water was from 9 to 50% of the outlet water. The number of zooplankton recruited into the pond from the inlet water, after the prawns were stocked, was negligible and contributed little to changes observed in zooplankton assemblages. Reproduction of barnacles within the pond appeared to play the most important role in changes in the assemblage. Water exchange did, however, appear to play a greater role in the changes observed in epibenthic fauna assemblages. In the last season of sampling the feeding of the dominant epibenthic species, Acetes sibogae, was examined using a combination of gut content and stable isotope analysis. Acetes gained little nutrition directly from the pelleted feed, probably relying primarily on zooplankton as their direct food source. Other dietary items such as macroalgae also played a role in the nutrition of the Acetes. If Acetes numbers were high at the beginning of a season they may compete with the newly stocked prawns for the zooplankton resource. However, they will not compete with the prawns later in the season when the prawns are gaining most of their nutrition from the pelleted feed. Overall it appears that zooplankton are important to the nutrition of the prawns at the beginning of the season when the assemblage is usually dominated by copepods. Later in the season the assemblage is dominated by barnacle nauplii which are recruited from within the pond. The establishment of an abundant assemblage of suitable zooplankton species before stocking prawn postlarvae would appear to be beneficial, if not essential. The assemblage of epibenthic fauna changes throughout the season as new recruits are brought in from outside the pond. Epibenthic faunal assemblages in ponds from southeast Queensland are dominated by Acetes which are not likely to adversely affect the production of prawns unless they are particularly abundant early in the grow out season when the prawns would be utilising the same food resources as Acetes.

Background: The loss of photosynthesis has occurred often in eukaryotic evolution, even more than its acquisition, which occurred at least nine times independently and which generated the evolution of the supergroups Archaeplastida, Rhizaria, Chromalveolata and Excavata. This secondary loss of autotrophic capability is essential to explain the evolution of eukaryotes and the high diversity of protists, which has been severely underestimated until recently. However, the ecological and evolutionary scenarios behind this evolutionary ‘‘step back’’ are still largely unknown. Methodology/Principal Findings: Using a dynamic model of heterotrophic and mixotrophic flagellates and two types of prey, large bacteria and ultramicrobacteria, we examine the influence of DOC concentration, mixotroph’s photosynthetic growth rate, and external limitations of photosynthesis on the coexistence of both types of flagellates. Our key premises are: large bacteria grow faster than small ones at high DOC concentrations, and vice versa; and heterotrophic flagellates are more efficient than the mixotrophs grazing small bacteria (both empirically supported). We show that differential efficiency in bacteria grazing, which strongly depends on cell size, is a key factor to explain the loss of photosynthesis in mixotrophs (which combine photosynthesis and bacterivory) leading to purely heterotrophic lineages. Further, we show in what conditions an heterotroph mutant can coexist, or even out-compete, its mixotrophic ancestor, suggesting that bacterivory and cell size reduction may have been major triggers for the diversification of eukaryotes. Conclusions/Significance: Our results suggest that, provided the mixotroph’s photosynthetic advantage is not too large, the (small) heterotroph will also dominate in nutrient-poor environments and will readily invade a community of mixotrophs and bacteria, due to its higher efficiency exploiting the ultramicrobacteria. As carbon-limited conditions were presumably widespread throughout Earth history, such a scenario may explain the numerous transitions from phototrophy to mixotrophy and further to heterotrophy within virtually all major algal lineages. We challenge prevailing concepts that affiliated the evolution of phagotrophy with eutrophic or strongly light-limited environments only.