For a complete picture of the lake ecosystem, it is necessary to understand the mechanisms regulating biomass and nutrient status (nutrient limitation, optimal supply, or surplus) of benthic algae, which are important primary producers and a food resource for grazers. This thesis gives an overview of the natural variation of benthic algae at different scales of space and time and on different substrates, and unravels some of the underlying factors. Algal nutrient status was assessed using the C:N:P (carbon:nitrogen:phosphorus) ratio of the entire natural benthic community. A review, observations, and experiments confirmed that a C:N:P ratio of about 158:18:1 (molar basis) represented an optimal nutrient supply, and that substantially higher C:N, N:P, or C:P ratios reflected algal growth limitation caused by an N or P nutrient deficiency. Horizontal variation of benthic algal biomass and nutrient status was patchy, of similar amount for all investigated distances, substrates, and lakes, and constituted a dominant proportion of the total variation. For example, patches of nutrient limited algae were found within only 10 m distance from patches with a nutrient surplus. Thus, horizontal variation should not be neglected when sampling benthic algae in lakes. Field observations suggested an impact of wind, nutrients, and grazers on the horizontal variation. Light and nutrients might have caused the observed vertical and temporal variation. Field experiments confirmed a simultaneous control of benthic algal biomass by nutrients and grazing, mediated by light and temperature. Grazing effects were larger than nutrient effects, but the comparison of natural communities in lakes of different trophy suggested that benthic algal biomass was controlled by nutrients in the long run. An important nutrient supply was animal excretions, causing a low C:N:P ratio of epizoon on zebra mussels, and algal communities associated with macrograzers. A field experiment revealed that 15N circulated one week longer in epizoon associated with a sessile caddisfly than in surrounding epilithon. In conclusion, the regulation of benthic algal biomass and nutrient status in lakes is complex, and benthic animals should be looked at not only as grazers, but also as a nutrient source.

No description available.

No description available.

Central Appalachian headwater streams in coalfield areas are prone to mining disturbances, and compensatory mitigation is required in cases of documented impacts. Stream construction on reclaimed mines is a common mitigation strategy. Streams constructed as compensatory mitigation are meant to restore structural and functional attributes of headwater streams and are often evaluated by measuring structural ecosystem characteristics. However, replacement of stream ecosystem functions is essential for mitigation of mining disturbances from an ecosystem perspective. This research compared selected structural and functional measures in eight constructed streams on mined areas to those of four forested reference streams across two years. Three organic matter functions were evaluated: riparian litterfall input, leaf breakdown, and periphyton accrual. Constructed streams were typically warmer than reference streams and also had elevated specific conductance, elevated oxidized nitrogen concentrations, depressed benthic macroinvertebrate richness, and lower levels of canopy cover. Functionally, litterfall input and total leaf breakdown means for constructed streams were approximately 25% and 60% of reference means, respectively. Leaf breakdown in constructed streams appeared to be inhibited as a result of reduced processing by benthic macroinvertebrates as well as inhibition of microbial and physicochemical pathways. Constructed streams with total breakdown rates most similar to reference-stream levels had the coldest stream temperatures. Areal periphyton biomass, benthic algal standing crop, and senescent autotrophic organic matter in constructed streams were roughly quadruple, double, and quintuple those of reference streams, respectively. Indicator ratios also suggested stream-type differences in periphyton structure. Mean algal accrual was greater in constructed streams than in reference streams during leaf-on seasons. My results suggest that light is likely the primary factor driving accrual rate differences during summer and fall, but that temperature may also be important during fall. Planting a diverse assemblage of native riparian trees and ensuring their successful development can inhibit benthic irradiance and thermal energy inputs while providing similar quantity and quality of OM to constructed streams, thereby fostering replacement of reference-like OM functions in some streams. / Ph. D.

Cyanobacteria can form nuisance proliferations and produce large concentrations of toxins that pose a health hazard. This thesis investigates cyanotoxin production by New Zealand benthic cyanobacteria. Cyanobacteria were sampled from lakes, reservoirs, streams, and rivers. Thirty-five strains were isolated into culture and screened for genes involved in the biosynthesis of common cyanotoxins. Positive results were confirmed and cyanotoxin concentrations quantified using analytical chemistry techniques. Genes involved in anatoxin a/homoanatoxin a biosynthesis were detected in nine out of ten Phormidium cf. uncinatum strains isolated from a single mat. Anatoxin a was confirmed in these strains by LC–MS/MS at concentrations from 0.3 to 6.4 mg kg⁻¹. One strain also produced homoanatoxin-a. Anatoxin-a variation between strains may explain the wide range in anatoxin a concentrations previously observed in New Zealand. The sxtA gene involved in saxitoxin biosynthesis was identified in Scytonema cf. crispum strains. Saxitoxin was confirmed in strains and environmental samples by Jellett PSP Rapid Test and HPLC–FD. Gonyautoxins, neosaxitoxin, and decarbamoyl derivatives were also detected. This study is the first identification of these compounds in Scytonema and in New Zealand cyanobacterial strains. These strains were isolated from recreational and pre-treatment drinking water reservoirs, highlighting the risk benthic cyanobacteria pose to human and animal health. Experiments were undertaken using cultures of Phormidium and Scytonema to determine how growth influences cyanotoxin production. The effects of iron and copper stress on P. autumnale were also investigated. High iron concentrations disrupted attachment mechanisms. Iron and copper had a significant effect on growth, without significantly affecting anatoxin a production. However, the maximum anatoxin a quota was consistently observed during early exponential growth. Scytonema cf. crispum produced higher saxitoxin quota throughout exponential growth than during the stationary phase. Both the Phormidium and Scytonema growth experiments indicate that high toxin quota can be expected early in benthic mat development, making early detection of these proliferations important.

Os estudos apresentados nesta tese tiveram como objetivo compreender aspectos da organização de assembleias de algas bênticas em riachos. Foi avaliado especificamente o efeito da heterogeneidade de habitat na escala de rugosidade de substratos e sua interação com distúrbios hidrológicos e organismos pastejadores (grazers). Utilizou-se uma abordagem experimental in situ com substratos lisos e rugosos (com fendas) constituindo os tratamentos. Os experimentos foram conduzidos no Rio do Marco, um riacho de quarta ordem na região dos Campos de Cima da Serra, em São José dos Ausentes, Rio Grande do Sul. Após uma introdução geral acerca do assunto são apresentados os capítulos com os estudos. O Capítulo 1 avaliou os efeitos da rugosidade de substratos sobre a riqueza de espécies, densidade, composição e padrões de distribuição de assembleias de algas bênticas. Assembleias em substratos rugosos foram mais ricas que assembleias em substratos lisos, porém esse resultado diferiu entre grupos com diferentes formas de vida, assim como as diferenças na composição de espécies, indicando que somente alguns grupos foram beneficiados pela ocorrência de refúgios em substratos rugosos. Observou-se também que diferenças na composição da assembleia entre substratos lisos e rugosos resultaram de aninhamento e substituição de espécies. No Capítulo 2 foi testada a hipótese de que substratos rugosos abrigam assembleias mais persistentes que substratos lisos. Os resultados encontrados confirmaram esta hipótese e foram atribuídos ao efeito da rugosidade per se após a exclusão de alguns fatores de confundimento, como estabilidade física dos substratos e diferenças na riqueza de espécies. Concluiu-se que o mecanismo mais plausível para explicar a maior persistência em substratos rugosos é a maior disponibilidade de refúgios em relação a substratos lisos. O objetivo do Capítulo 3 foi avaliar o efeito da rugosidade de substratos sobre assembleias de algas frente a um distúrbio hidrológico experimental, testando especificamente se a resistência e a resiliência de algas é maior em substratos rugosos que em substratos lisos e se a resposta de diferentes formas de vida é distinta. Os resultados indicaram que a rugosidade de substratos não apresenta efeitos pronunciados sobre a resistência e a resiliência de algas bênticas. Porém, grupos de algas com distintas formas de vida diferiram quanto à resistência e resiliência, de forma que o padrão observado está relacionado a atributos biológicos que conferem habilidades para suportar distúrbios. Os resultados deste estudo em conjunto com os resultados obtidos no Capítulo 2 e resultados obtidos por outros autores permitiram que fosse proposto que a importância da heterogeneidade de habitas e de refúgios é mediada pela intensidade de distúrbios. No Capítulo 4 foram avaliados os efeitos independentes e as interações de rugosidade de substratos, peixes pastejadores e tipos de mesohabitats (corredeiras e remansos) sobre a biomassa algal, matéria orgânica e peso seco total. O efeito de peixes pastejadores não foi mediado pelos mesohabitats, sendo que a exclusão de peixes causou aumento nas três variáveis resposta, independente do tipo de mesohabitat. Por outro lado, houve uma interação entre mesohabitats e substratos para determinar o acúmulo de peso seco total, sendo que substratos rugosos acumularam maior quantidade de material em remansos que em corredeiras, enquanto substratos lisos acumularam quantidades similares entre os dois tipos de mesohabitats. Já a biomassa algal e matéria orgânica apresentaram maior acúmulo nos substratos rugosos em relação aos lisos, independente do mesohabitat. Os resultados encontrados indicam que a perda de organismos pastejadores pode afetar o acúmulo de biomassa algal e material orgânico, importantes recursos alimentares em riachos. Finalmente, na última seção são apresentadas as considerações finais. / The studies presented in this thesis aimed at understanding some aspects of the organization of benthic algal assemblages in streams. Specifically, the studies evaluated the effect of habitat heterogeneity on the scale of substrate roughness and its interaction with hydrological disturbances and grazing organisms. An experimental in situ approach was used, in which the treatments were constituted by smooth and rough (with crevices) substrates. The experiments were conducted in the Marco River, a fourth-order stream at the region of Campos de Cima da Serra, São José dos Ausentes, Rio Grande do Sul, Brazil. After a general introduction on the subjects of this thesis, the chapters with the studies are presented. The Chapter 1 describes a study to evaluate the effects of substrate roughness on species richness, density, composition and distribution patterns of benthic algal assemblages. Algal assemblages on rough substrates were richer than on smooth substrates, but this result differed among algal life forms. Similarly, results on species composition also differed among life forms, indicating that only some groups were benefited by the occurrence of refuges on rough substrates. It was also observed that differences in the composition of assemblages between rough and smooth substrates resulted from nestedness and turnover. The Chapter 2 describes a study in which was tested the hypothesis that rough substrates harbor more persistent assemblages than smooth substrates. The results confirmed this hypothesis and were attributed to the effect of roughness per se after the exclusion of some confounding factors, such as the physical stability of substrates and differences in species richness. It was concluded that the most plausible mechanism that could explain the higher persistence in rough substrates is the greater availability of refuges in relation to smooth substrates. The main aim of the study described in Chapter 3 was to evaluate the effect of substrate roughness on algal assemblages in the face of an experimental hydrological disturbance. Specifically, it was tested if the resistance and resilience of algae is higher on rough than on smooth substrates and if algal life forms differ in their response to disturbance. The results indicated that substrate roughness had no pronounced effects on benthic algal resistance and resilience. However, algal life forms differ in their resistance and resilience, and the observed pattern was mostly related to biological traits that confer abilities to support disturbances. The results of this study, in addition to the results obtained in Chapters 1 and 2, allowed us to suggest that the importance of habitat heterogeneity and refuges is mediated by the intensity of disturbances. The Chapter 4 describes a study that evaluated the independent and interacting effects of substrate roughness, grazing fish and mesohabitats (pools and riffles) on algal biomass, organic matter and total dry mass accrual. The effect of grazers was not mediated by mesohabitats, and their exclusion caused an increase in the three response variables, regardless of mesohabitat. On the other hand, mesohabitats and substrates interacted to determine the accrual of total dry mass, since rough substrates accumulated greater amounts of dry mass in pools than in riffles, while smooth substrates accumulated similar amounts in both mesohabitats. Algal biomass and organic matter showed greater accrual on rough than on smooth substrates, independently of mesohabitat. These results indicate that the loss of grazers may affect the accrual of algal biomass and organic matter, which are important food resources in streams. Finally, in the last section are presented the concluding remarks.

Os estudos apresentados nesta tese tiveram como objetivo compreender aspectos da organização de assembleias de algas bênticas em riachos. Foi avaliado especificamente o efeito da heterogeneidade de habitat na escala de rugosidade de substratos e sua interação com distúrbios hidrológicos e organismos pastejadores (grazers). Utilizou-se uma abordagem experimental in situ com substratos lisos e rugosos (com fendas) constituindo os tratamentos. Os experimentos foram conduzidos no Rio do Marco, um riacho de quarta ordem na região dos Campos de Cima da Serra, em São José dos Ausentes, Rio Grande do Sul. Após uma introdução geral acerca do assunto são apresentados os capítulos com os estudos. O Capítulo 1 avaliou os efeitos da rugosidade de substratos sobre a riqueza de espécies, densidade, composição e padrões de distribuição de assembleias de algas bênticas. Assembleias em substratos rugosos foram mais ricas que assembleias em substratos lisos, porém esse resultado diferiu entre grupos com diferentes formas de vida, assim como as diferenças na composição de espécies, indicando que somente alguns grupos foram beneficiados pela ocorrência de refúgios em substratos rugosos. Observou-se também que diferenças na composição da assembleia entre substratos lisos e rugosos resultaram de aninhamento e substituição de espécies. No Capítulo 2 foi testada a hipótese de que substratos rugosos abrigam assembleias mais persistentes que substratos lisos. Os resultados encontrados confirmaram esta hipótese e foram atribuídos ao efeito da rugosidade per se após a exclusão de alguns fatores de confundimento, como estabilidade física dos substratos e diferenças na riqueza de espécies. Concluiu-se que o mecanismo mais plausível para explicar a maior persistência em substratos rugosos é a maior disponibilidade de refúgios em relação a substratos lisos. O objetivo do Capítulo 3 foi avaliar o efeito da rugosidade de substratos sobre assembleias de algas frente a um distúrbio hidrológico experimental, testando especificamente se a resistência e a resiliência de algas é maior em substratos rugosos que em substratos lisos e se a resposta de diferentes formas de vida é distinta. Os resultados indicaram que a rugosidade de substratos não apresenta efeitos pronunciados sobre a resistência e a resiliência de algas bênticas. Porém, grupos de algas com distintas formas de vida diferiram quanto à resistência e resiliência, de forma que o padrão observado está relacionado a atributos biológicos que conferem habilidades para suportar distúrbios. Os resultados deste estudo em conjunto com os resultados obtidos no Capítulo 2 e resultados obtidos por outros autores permitiram que fosse proposto que a importância da heterogeneidade de habitas e de refúgios é mediada pela intensidade de distúrbios. No Capítulo 4 foram avaliados os efeitos independentes e as interações de rugosidade de substratos, peixes pastejadores e tipos de mesohabitats (corredeiras e remansos) sobre a biomassa algal, matéria orgânica e peso seco total. O efeito de peixes pastejadores não foi mediado pelos mesohabitats, sendo que a exclusão de peixes causou aumento nas três variáveis resposta, independente do tipo de mesohabitat. Por outro lado, houve uma interação entre mesohabitats e substratos para determinar o acúmulo de peso seco total, sendo que substratos rugosos acumularam maior quantidade de material em remansos que em corredeiras, enquanto substratos lisos acumularam quantidades similares entre os dois tipos de mesohabitats. Já a biomassa algal e matéria orgânica apresentaram maior acúmulo nos substratos rugosos em relação aos lisos, independente do mesohabitat. Os resultados encontrados indicam que a perda de organismos pastejadores pode afetar o acúmulo de biomassa algal e material orgânico, importantes recursos alimentares em riachos. Finalmente, na última seção são apresentadas as considerações finais. / The studies presented in this thesis aimed at understanding some aspects of the organization of benthic algal assemblages in streams. Specifically, the studies evaluated the effect of habitat heterogeneity on the scale of substrate roughness and its interaction with hydrological disturbances and grazing organisms. An experimental in situ approach was used, in which the treatments were constituted by smooth and rough (with crevices) substrates. The experiments were conducted in the Marco River, a fourth-order stream at the region of Campos de Cima da Serra, São José dos Ausentes, Rio Grande do Sul, Brazil. After a general introduction on the subjects of this thesis, the chapters with the studies are presented. The Chapter 1 describes a study to evaluate the effects of substrate roughness on species richness, density, composition and distribution patterns of benthic algal assemblages. Algal assemblages on rough substrates were richer than on smooth substrates, but this result differed among algal life forms. Similarly, results on species composition also differed among life forms, indicating that only some groups were benefited by the occurrence of refuges on rough substrates. It was also observed that differences in the composition of assemblages between rough and smooth substrates resulted from nestedness and turnover. The Chapter 2 describes a study in which was tested the hypothesis that rough substrates harbor more persistent assemblages than smooth substrates. The results confirmed this hypothesis and were attributed to the effect of roughness per se after the exclusion of some confounding factors, such as the physical stability of substrates and differences in species richness. It was concluded that the most plausible mechanism that could explain the higher persistence in rough substrates is the greater availability of refuges in relation to smooth substrates. The main aim of the study described in Chapter 3 was to evaluate the effect of substrate roughness on algal assemblages in the face of an experimental hydrological disturbance. Specifically, it was tested if the resistance and resilience of algae is higher on rough than on smooth substrates and if algal life forms differ in their response to disturbance. The results indicated that substrate roughness had no pronounced effects on benthic algal resistance and resilience. However, algal life forms differ in their resistance and resilience, and the observed pattern was mostly related to biological traits that confer abilities to support disturbances. The results of this study, in addition to the results obtained in Chapters 1 and 2, allowed us to suggest that the importance of habitat heterogeneity and refuges is mediated by the intensity of disturbances. The Chapter 4 describes a study that evaluated the independent and interacting effects of substrate roughness, grazing fish and mesohabitats (pools and riffles) on algal biomass, organic matter and total dry mass accrual. The effect of grazers was not mediated by mesohabitats, and their exclusion caused an increase in the three response variables, regardless of mesohabitat. On the other hand, mesohabitats and substrates interacted to determine the accrual of total dry mass, since rough substrates accumulated greater amounts of dry mass in pools than in riffles, while smooth substrates accumulated similar amounts in both mesohabitats. Algal biomass and organic matter showed greater accrual on rough than on smooth substrates, independently of mesohabitat. These results indicate that the loss of grazers may affect the accrual of algal biomass and organic matter, which are important food resources in streams. Finally, in the last section are presented the concluding remarks.

Os estudos apresentados nesta tese tiveram como objetivo compreender aspectos da organização de assembleias de algas bênticas em riachos. Foi avaliado especificamente o efeito da heterogeneidade de habitat na escala de rugosidade de substratos e sua interação com distúrbios hidrológicos e organismos pastejadores (grazers). Utilizou-se uma abordagem experimental in situ com substratos lisos e rugosos (com fendas) constituindo os tratamentos. Os experimentos foram conduzidos no Rio do Marco, um riacho de quarta ordem na região dos Campos de Cima da Serra, em São José dos Ausentes, Rio Grande do Sul. Após uma introdução geral acerca do assunto são apresentados os capítulos com os estudos. O Capítulo 1 avaliou os efeitos da rugosidade de substratos sobre a riqueza de espécies, densidade, composição e padrões de distribuição de assembleias de algas bênticas. Assembleias em substratos rugosos foram mais ricas que assembleias em substratos lisos, porém esse resultado diferiu entre grupos com diferentes formas de vida, assim como as diferenças na composição de espécies, indicando que somente alguns grupos foram beneficiados pela ocorrência de refúgios em substratos rugosos. Observou-se também que diferenças na composição da assembleia entre substratos lisos e rugosos resultaram de aninhamento e substituição de espécies. No Capítulo 2 foi testada a hipótese de que substratos rugosos abrigam assembleias mais persistentes que substratos lisos. Os resultados encontrados confirmaram esta hipótese e foram atribuídos ao efeito da rugosidade per se após a exclusão de alguns fatores de confundimento, como estabilidade física dos substratos e diferenças na riqueza de espécies. Concluiu-se que o mecanismo mais plausível para explicar a maior persistência em substratos rugosos é a maior disponibilidade de refúgios em relação a substratos lisos. O objetivo do Capítulo 3 foi avaliar o efeito da rugosidade de substratos sobre assembleias de algas frente a um distúrbio hidrológico experimental, testando especificamente se a resistência e a resiliência de algas é maior em substratos rugosos que em substratos lisos e se a resposta de diferentes formas de vida é distinta. Os resultados indicaram que a rugosidade de substratos não apresenta efeitos pronunciados sobre a resistência e a resiliência de algas bênticas. Porém, grupos de algas com distintas formas de vida diferiram quanto à resistência e resiliência, de forma que o padrão observado está relacionado a atributos biológicos que conferem habilidades para suportar distúrbios. Os resultados deste estudo em conjunto com os resultados obtidos no Capítulo 2 e resultados obtidos por outros autores permitiram que fosse proposto que a importância da heterogeneidade de habitas e de refúgios é mediada pela intensidade de distúrbios. No Capítulo 4 foram avaliados os efeitos independentes e as interações de rugosidade de substratos, peixes pastejadores e tipos de mesohabitats (corredeiras e remansos) sobre a biomassa algal, matéria orgânica e peso seco total. O efeito de peixes pastejadores não foi mediado pelos mesohabitats, sendo que a exclusão de peixes causou aumento nas três variáveis resposta, independente do tipo de mesohabitat. Por outro lado, houve uma interação entre mesohabitats e substratos para determinar o acúmulo de peso seco total, sendo que substratos rugosos acumularam maior quantidade de material em remansos que em corredeiras, enquanto substratos lisos acumularam quantidades similares entre os dois tipos de mesohabitats. Já a biomassa algal e matéria orgânica apresentaram maior acúmulo nos substratos rugosos em relação aos lisos, independente do mesohabitat. Os resultados encontrados indicam que a perda de organismos pastejadores pode afetar o acúmulo de biomassa algal e material orgânico, importantes recursos alimentares em riachos. Finalmente, na última seção são apresentadas as considerações finais. / The studies presented in this thesis aimed at understanding some aspects of the organization of benthic algal assemblages in streams. Specifically, the studies evaluated the effect of habitat heterogeneity on the scale of substrate roughness and its interaction with hydrological disturbances and grazing organisms. An experimental in situ approach was used, in which the treatments were constituted by smooth and rough (with crevices) substrates. The experiments were conducted in the Marco River, a fourth-order stream at the region of Campos de Cima da Serra, São José dos Ausentes, Rio Grande do Sul, Brazil. After a general introduction on the subjects of this thesis, the chapters with the studies are presented. The Chapter 1 describes a study to evaluate the effects of substrate roughness on species richness, density, composition and distribution patterns of benthic algal assemblages. Algal assemblages on rough substrates were richer than on smooth substrates, but this result differed among algal life forms. Similarly, results on species composition also differed among life forms, indicating that only some groups were benefited by the occurrence of refuges on rough substrates. It was also observed that differences in the composition of assemblages between rough and smooth substrates resulted from nestedness and turnover. The Chapter 2 describes a study in which was tested the hypothesis that rough substrates harbor more persistent assemblages than smooth substrates. The results confirmed this hypothesis and were attributed to the effect of roughness per se after the exclusion of some confounding factors, such as the physical stability of substrates and differences in species richness. It was concluded that the most plausible mechanism that could explain the higher persistence in rough substrates is the greater availability of refuges in relation to smooth substrates. The main aim of the study described in Chapter 3 was to evaluate the effect of substrate roughness on algal assemblages in the face of an experimental hydrological disturbance. Specifically, it was tested if the resistance and resilience of algae is higher on rough than on smooth substrates and if algal life forms differ in their response to disturbance. The results indicated that substrate roughness had no pronounced effects on benthic algal resistance and resilience. However, algal life forms differ in their resistance and resilience, and the observed pattern was mostly related to biological traits that confer abilities to support disturbances. The results of this study, in addition to the results obtained in Chapters 1 and 2, allowed us to suggest that the importance of habitat heterogeneity and refuges is mediated by the intensity of disturbances. The Chapter 4 describes a study that evaluated the independent and interacting effects of substrate roughness, grazing fish and mesohabitats (pools and riffles) on algal biomass, organic matter and total dry mass accrual. The effect of grazers was not mediated by mesohabitats, and their exclusion caused an increase in the three response variables, regardless of mesohabitat. On the other hand, mesohabitats and substrates interacted to determine the accrual of total dry mass, since rough substrates accumulated greater amounts of dry mass in pools than in riffles, while smooth substrates accumulated similar amounts in both mesohabitats. Algal biomass and organic matter showed greater accrual on rough than on smooth substrates, independently of mesohabitat. These results indicate that the loss of grazers may affect the accrual of algal biomass and organic matter, which are important food resources in streams. Finally, in the last section are presented the concluding remarks.

The role of communities associated to a substrate as fundamental components of an aquatic system has been recognized by many authors. Although periphyton has been considered of great importance in shallow lakes, its inclusion in different ecological models has been unusual. Phillips et al. (1978) were the first in considering epiphyton as an internal mechanism leading the system to a change in its state from greater water transparecy to more turbid conditions. In contrast, Scheffer et al. (1993) proposed the Alternative Equilibria Hypothesis (AEH), which considers that a shallow lake can alternate between two states, a Clear Water State (stabilized by the presence of submerged macrophytes) and a Turbid Water State (stabilized by an elevated phytoplankton biomass), excluding in this scheme the periphytic communities. Later only the conceptual model coined by Goldsborough and Robinson (1996, G&R model) emphasizes again the importance of epiphyton as well as that of epipelon and metaphyton by establishing the possibility of occurrence of four states according to the contributions to the total algal biomass of each of the algal taxocenosis. Some analyses of the system’s dynamics have been performed in pampean lakes in the context of the AEH. On the other hand, knowledge regarding periphytic and benthic communities is still poor. We propose as an hypothesis that phytoperiphyton will respond to the occurrence of different equilibria states, displaying, at the same time, strong interrelationships with the phytobenthos as well as with the phytoplankton. On the other hand, we consider the possibility that the phytoperiphytic and phytobenthic assemblages may function as internal mechanisms both, as state change or as state stability factors. Finally, we assess the importance of phytoperiphyton as regards its contributions to the total algal biomass in the context of the G&R model, comparing then the results with those obtained in the case of the AEH. The general objectives of the present investigation are to gather information about the structure and dynamics of the phytoperiphyton and its relationships with environmental factors as well as with the phytobenthon and phytoplankton in a pampean shallow lake; and to analyze phytoperiphytic dynamics under the AEH and the G&R model contexts, evaluating periphytic contributions to the total algal biomass. This investigation was carried out in an seepage shallow lake, Lacombe Lake, which is included in the “Grupo Marginal de lagunas” group in the pericoastal region of Provincia de Buenos Aires. Sampling was performed from July 2001 to June 2002 at four sites according to their exposure to environmental factors and the presence of macrophytes: bulrush beds, (JP and JC); and sites devoided of emergent vegetation but with stands of submerged macrophytes during the growing season (ALe and ALr). J sites and ALe were also representative of deeper zones and ALr of a site near the shore. Environmental variables were measured in situ and water samples were collected for chemical analysis and for plankton identification and quantification, following a vertical profile (excepting in ALr). In the case of phytoperiphyton, bulrush epiphyte samples (epiphyton) were taken following a vertical profile and those of submerged macrophytes were collected from the first 25 cm of the stems. Benthon (epipelon) was sampled using a corer. Aliquots for chemical analysis were also extracted from those samples. Vegetal substrate dynamics was also evaluated during sampling. At the laboratory, concentrations of algal nutrients, of the eight principal ions and of soluble polyphenols, as well as alkalinity and total hardness were measured. Phytoplanktonic, epiphytic and epipelic samples were treated for active a chlorophyll and pheopigment determinations. Organism identification and quantification were made according to standardized and own developed methods. First, variations in biotic and abiotic variables were analyzed for the determination of different equilibrium states occurrence. Three perturbation events were registered: precipitations in August, October and March. All of them involved water level increases and conductivity diminutions. Especially in the first two events, nutrient input from runoff was observed; meanwhile in March the input of water determined the dilution in nutrients concentrations. On the other hand, light conditions improved, particularly from October onwards, when stands of submerged macrophytes developed and a decrease of phytoplankton density and a change in its specific composition was detected. The latter being generated by the dilution effects of water input, the shading of the aquatic plants and algal consumption by cladocerans. March event also involved an improvement of light underwater climate but the increase in the water level affected macrophyte stands which mainly withdrawed to the shallowest zones. As regards phytoplankton, changes in the specific composition were recorded again. The statistic evaluation of these facts in conjunction with the analysis of permanence, resiliency and resistance properties of the dominating taxocenosis leads to interpret in the context of the AEH that two equilibria states occurred in the shallow lake. During the first three months a Turbid Water State developed, given by the phytoplankton domination given by an oscilatorial cyanophyte, minor transparency and relatively higher nutrient concentrations. In October and November the passage between states occurred, thus considering these months as transitional. From December onwards a Clear Water State established given by extended development of stands of submerged macrophytes, greater water transparency and minor nutrient concentrations. Finally, from March to May, given the submerged macrophyte withdrawal to shallowest sectors it can be interpreted that the system oscillated around a clear water state, or else, taking into account changes in the planktonic community, it is suggested that a third different state established, the “mixed phytoplankton” state. It is also suggested that states developed in Lacombe could be alternative. Next, the structure and dynamics of phytoperiphyton on bulrush and submerged macrophytes was analyzed. Vertical spatial and temporal variations of bulrush epiphyton showed to be more important than differences between sites. These variations involved smaller biomass values and a simpler physiognomy during turbid water phase and along the vertical gradient in all sampling occasions. In October, related to better light underwater conditions and the nutrient input, epiphytic biomass increased, principally promoted by species typical of the turbid phase. Nonetheless, a structural change started in this month, driven by herbivores but also by the modification of internal relationships between members of the same taxocenosis (shading and less bare substrate to be colonized). This resulted in the establishment of the characteristic assemblages of the clear water phase. During the first three months domination by stalked diatoms structured the community in a unique stratum, resistant to the less favourable light conditions and resilient to August perturbations. From December onwards Spirogyra species dominated, constituting the upper stratum, followed by Oedogonium species in the middle stratum and diatoms in the lower one. These assemblages were stabilized by mechanisms mainly displayed by the zignematalean taxon (shading, epifitism inhibition), allowing to considerer that they were resilient to March perturbation event (de novo establishment on non previously colonized surfaces). In face of the permanence, resistance and resilience properties showed by the taxocenosis, two different states of bulrush phytoperiphyton are considered: turbid water and clear water assemblages. This temporal pattern appeared less well delimited with depth and showed up later in time. Epiphyton data suggested that both states could be alternative. Differences between sites could be observed in respect to phytoperiphyton growing on submerged macrophytes, but the same temporal pattern showed by bulrush periphyton couldn’t be detected due to a great variability in the data gathered. In the next section, the analysis of the structure and dynamics of the phytobenton was carried out. It was showed that differences between the deepest sampling points and the shallowest site were greater than temporal ones. The composition of the assemblages was scarcely variable in time. The taxocenosis was principally related to the poor light conditions recorded on the sediment surface throughout the entire sampling period, even during the clear water phase. The examination of the specific composition in function of its origin evidenced the influence that the epipelon receives from other taxocenosis, constituting a refuge site for allochthonous taxa during unfavourable situations. The scarce variability of the epipelic fraction is also showed. In this way, it is considered that the taxocenosis maintained itself in only one state, indifferent to state changes in the lake and principally resisting the darkness and bearing to live in the unstable sediments. Subsequently, first the roles of the phytoperiphyton and complementary those of phytobenton, as state change or state stability agents were evaluated. The harmful effect of the phytoperiphyton on submerged macrophytes was probably counteracted by plant ramification production and allelopathy. The evidence suggested that its role as nutrient sequestrator is of minor importance in this case, due to other factors that might have disturbed phytoplankton more intensely than competition with phytoperiphyton. Much evidence supported the role of phytobenton as a source of epiphytic and planktonic species belonging either to turbid and clear water state assemblages. The results did not support the hypothesis that phytobenton may play a role as a sediment-stabilizing factor. On the contrary, it may be perturbating agent. Next, an analysis of the contributions of each community to the total algal biomass in the context of the G&R model was performed. Comparisons with AEH results were also carried out. This leaded to the consideration of the occurrence of the following states in Lacombe Lake: a Lake State consistent with the Turbid Water phase, an Open State consistent with the Clear Water phase and local Dry States in ALr site occurred both during the turbid water phase (July) or the clear water situation (March-April). The substantial contribution of epiphyton especially, that growing on submerged macrophytes, is emphasized. The importance of the discrimination of different fractions of epipelon material according to its origin when evaluating the contributions of each community to total algal biomass is also highlighted. Finally, a new interpretation is presented based on algal taxocenosis dynamics, which incorporates elements from both, the AEH and the G&R model. In conclusion, phytoperiphyton in this pampean shallow lake is strongly affected by the factors that determine the establishment or passage from one equilibrium state to another. These assemblages respond to changes through modifications in their own internal structure. These structures gradually alter with depth, remaining these changes more fully explained by the comparison with phytobenton dynamics. Besides the possible roles that the taxocenosis might undertake as an internal mechanism in the change or stabilization of systems states in different situations other than those encountered during this study, epiphyton importance in the systems context, is principally evidenced by the great contribution that these taxocenosis make to the total algal biomass, constituting the dominant compartment during clear water states. / El rol de las comunidades relacionadas a un sustrato como componentes fundamentales dentro de un sistema acuático ha sido reconocido por varios autores. A pesar de la importancia dada al perifiton en lagos someros, su inclusión dentro de diferentes modelos ecológicos ha sido inusual. Phillips et al. (1978) son los primeros en considerar al epifiton como un mecanismo interno en el pasaje de un estado de aguas transparentes a otro más turbio. En contraste, Scheffer et al. (1993), proponen la Hipótesis de Estados de Equilibrio Alternativos (HEEA) que considera que un lago somero puede alternar entre dos estados, uno de Aguas Claras, (estabilizado por la presencia de macrófitas sumegidas) y otro de Aguas Turbias (estabilizado por la elevada biomasa de fitoplancton), no tomando en consideración a las comunidades perifiticas. Sólo el marco conceptual propuesto por Goldsborough y Robinson (1996, modelo G&R) da nuevamente relevancia al epifiton, incluyendo al epipelon, al metafiton y al fitoplancton dentro de un modelo ecológico, discriminando cuatro estados de acuerdo a la dominancia en la biomasa algal total de alguna de estas taxocenosis. En las lagunas pampeanas se han realizado algunos análisis de la dinámica de los sistemas en el marco de la HEEA. Por otro lado, el conocimiento acerca de las comunidades perifíticas y bentónicas todavía es escaso. Se propone como hipótesis que el fitoperifiton variará de acuerdo a la ocurrencia de los diferentes estados en la laguna (según HEEA), interrelacionándose al mismo tiempo con el fitobenton y el fitoplancton. Por otro lado, también se considera la posibilidad de que tanto los ensambles fitoperifíticos como fitobentónicos sean mecanismos internos en el pasaje o en la estabilización de un estado. Por último, se valora la importancia del fitoperifiton en cuanto a sus aportes a la biomasa algal total en el marco del modelo G&R, realizando comparaciones con los resultados obtenidos en el caso de la HEEA. Los objetivos generales de esta investigación son: obtener información acerca de la dinámica y estructura del fitoperifiton y su relación con factores ambientales así como, también, acerca de sus interrelaciones con el fitobenton y el fitoplancton en una laguna pampeana; y analizar su dinámica en el marco de la HEEA y del modelo G&R valorando los aportes del fitoperifiton a la biomasa algal total. El estudio se realizó en la laguna Lacombe comprendida en el Grupo Marginal de lagunas en la región pericostera de la Provincia de Buenos Aires. Los muestreos se realizaron desde julio 2001 a junio 2002 en cuatro sitios de acuerdo a su grado de exposición a las variables ambientales y la presencia de macrófitas: Juncales (JP y JC); sitios de Aguas Libres con o sin macrófitas sumergidas durante la época de crecimiento (ALe y ALr). Los sitios J y ALe representan, puntos de mayor profundidad y ALr un sitio cercano a la orilla. Siguiendo un perfil vertical (con excepción del sitio cercano a la orilla, ALr) se midieron variables ambientales in situ y se tomaron muestras de agua para realizar análisis químicos del agua y la identificación y la cuantificación del plancton. En el caso del fitoperifiton se tomaron muestras de epifitos sobre junco (epifiton) también en un perfil vertical y en los 25 cm apicales del tallo de plantas sumergidas. El benton (epipelon) fue muestreado con un corer, extrayendo muestras también para realizar análisis químicos de los sedimentos. En el campo también se evaluaron las dinámicas de los sustratos vegetales. En el laboratorio se midieron las concentraciones de nutrientes algales, de los ocho iones principales, de los polifenoles solubles así como también la reserva alcalina y la dureza total. Las muestras de fitoplancton, epifiton y epipelon fueron tratadas para la estimación de clorofila a activa y feopigmentos, y la identificación y cuantificación de organismos según métodos estandarizados y propios. En principio, se analizaron las variables bióticas y abióticas para determinar la ocurrencia de estados de equilibrio en la laguna. Tres eventos principales de perturbación ocurrieron durante el período de estudio: las precipitaciones en agosto, octubre y marzo. Todas ellas involucraron aumentos del nivel hidrométrico y disminuciones en la conductividad. En los dos primeros, especialmente en octubre, se produjo la entrada de nutrientes por escorrentía, en tanto que en marzo el aporte de agua resultó en la dilución de los mismos. Por otro lado, las condiciones lumínicas mejoraron especialmente a partir de octubre, lo que se relacionó con el desarrollo de manchones de macrófitas sumergidas y una menor densidad y composición diferente del fitoplancton. Esto último se produjo por los efectos de dilución de la entrada de agua, el sombreado de las plantas acuáticas y por el pastoreo de cladóceros. El evento de marzo involucró también un mejoramiento de las condiciones de luz en la columna de agua pero, paralelamente, determinó la retracción de los manchones de macrófitas sumergidas hasta zonas más cercanas a la orilla. En el fitoplancton se operaron cambios en la composición específica. La evaluación estadística de estos hechos junto con el análisis de las características de permanencia, resiliencia y resistencia de las taxocenosis dominantes condujeron a interpretar que durante el período de estudio se establecen dos estados de equilibrio según la HEEA. Durante los tres primeros meses se desarrolló un estado de aguas turbias definido por la dominancia del fitoplancton con el predominio de una oscilatorial, menor transparencia y concentraciones de nutrientes relativamente más elevadas. En octubre y noviembre se produjo el pasaje entre estados, considerándose a estos meses como transicionales. De diciembre hasta el final del estudio, se desarrolló un estado de aguas claras definido por la presencia de manchones de macrófitas, mayor transparencia del agua y menores concentraciones de nutrientes. Finalmente, de marzo hasta mayo, dada la retracción de los manchones de macrófitas sumergidas, se puede interpretar que ocurrió una oscilación del sistema alrededor de un estado de aguas claras o, dados los cambios operados en el plancton, un tercer estado “mixed phytoplankton”. Se evidenció que los estados establecidos podrían ser alternativos. Seguidamente se analizaron la estructura y dinámica del fitoperifiton sobre junco y las macrófitas sumergidas. Las variaciones espaciales verticales y temporales en la dinámica del perifiton sobre junco resultaron más marcadas que las espaciales horizontales (entre sitios). Estas variaciones implicaron una menor biomasa y una fisonomía más simple durante la fase de aguas turbias y a lo largo del perfil vertical en todas las fechas. Relacionado con el incremento de la luz en la columna de agua y por la entrada de nutrientes en octubre, la biomasa epifítica aumentó, impulsada por las especies características de la fase turbia. Sin embargo, a partir de ese mes se dió un cambio estructural, generado por el pastoreo pero, también, por la modificación en las relaciones entre los componentes de la taxocenosis (autosombreado y menor disponibilidad de sustrato libre) que desembocó en el establecimiento de los ensambles característicos de la fase clara. Durante los tres primeros meses el predominio de diatomeas pedunculadas conformó una comunidad uniestrato, resistente a una menor disponibilidad de la luz y resiliente frente a las perturbaciones de agosto. De diciembre a febrero se desarrolló un ensamble donde predominan las especies de Spirogyra formando un estrato superior, las de Oedogonium en un estrato medio y las diatomeas en un estrato inferior. Estos ensambles presentaron mecanismos de estabilización dados por los talos de la zignematal (sombreado, inhibición del crecimiento de epifitos secundarios), considerándose resilientes frente a las perturbaciones de marzo (desarrollo de los ensambles en sitios previamente no colonizados). En función de las características de permanencia, resistencia y resiliencia, se diferenciaron dos estados en el fitoperifiton de junco: ensambles de fase turbia y ensambles de fase clara. Este patrón temporal se hizo menos marcado en profundidad y aparece en forma retrasada en el tiempo. Los datos habrían indicado la posibilidad de que ambos estados fuesen alternativos. El fitoperifiton sobre macrófitas sumergidas presentó variaciones definidas en el sentido espacial horizontal (mayor biomasa en sitios más profundos) pero no se detectó un patrón temporal, similar al del perifiton sobre junco, dada la gran variabilidad de los datos obtenidos. En el apartado siguiente se analizaron la estructura y dinámica del fitobenton. Se estableció que las variaciones entre sitios de muestreo fueron más marcadas que las temporales. Los ensambles epipélicos presentaron en general una composición escasamente variable a lo largo del tiempo relacionándose principalmente con la disponibilidad de luz. Ésta fue escasa durante todo el período de estudio por lo que las condiciones de crecimiento para las algas béntonicas no fueron favorables, aún con los cambios de estado en la laguna. El análisis del material en función de su origen evidenció las influencias que esta comunidad recibe desde otros ensambles, siendo un sitio de refugio para organismos alóctonos durante épocas desfavorables. También se evidenció la escasa variabilidad de la fracción epipélica propiamente dicha durante todo el período de estudio. Así, se considera que esta taxocenosis se mantuvo en un solo estado, ajena al pasaje de estado en la laguna, principalmente resistiendo la falta de luz y la inestabilidad de los sedimentos. Posteriormente, se evaluó primeramente el papel del fitoperifiton y, en forma complementaria, el del fitobenton como impulsores del pasaje de estados o como estabilizadores de los mismos en la laguna. El efecto de sombreado del fitoperifiton sobre macrófitas sumergidas probablemente fue contrarrestado por la producción de nuevas ramificaciones de las macrófitas y por alelopatía. Las evidencias señalaron que su rol como secuestradores de nutrientes disponibles para el fitoplancton fue menor en este caso, dado que éste último se halla perturbado por otros factores más que por la competencia con el epifiton. Las evidencias señalaron al epipelon como simiente de especies del epifiton y del fitoplancton tanto pertenecientes a la fase turbia como a la clara. En cambio, los resultados no avalaron su rol como estabilizador de los sedimentos. Al contrario, podría ser una perturbación de éstos. A continuación, se examinaron las contribuciones de las taxocenosis a la biomasa algal total en el marco del modelo G&R. También se realizó la comparación de los resultados con aquellos obtenidos a partir de la HEEA. Se consideró la ocurrencia de un estado Lake State congruente con la fase turbia en la laguna, un estado Open State congruente con la fase clara y estados locales Dry State en ALr tanto en la fase turbia (julio) como en la clara (marzo-abril). Se destacó la substancial contribución del epifiton sobre macrófitas sumergidas. También, se subrayó la importancia de discriminar, en relación con el epipelon, los aportes de cada una de las taxocenosis cuando se evalúan las contribuciones a la biomasa algal total. Finalmente, se ofreció una interpretación basada sobre la dinámica de las taxocenosis algales considerando elementos de la HEEA y del modelo G&R. En conclusión, el fitoperifiton en las lagunas pampeanas fue fuertemente influido por los factores que determinan los estados de equilibrio, respondiendo al cambio de los mismos con la modificación en su estructura interna. Ésta se alteró gradualmente con la profundidad, quedando estos cambios más completamente explicados a través de la comparación con la dinámica del fitobenton. Además de los posibles roles que podría desempeñar como mecanismo interno de pasaje o como estabilizador de estados en situaciones diferentes a las halladas en Lacombe, su importancia en el contexto del sistema se manifiesta en el gran aporte que realiza a la biomasa algal total, constituyendo el compartimiento dominante en los estados de aguas claras.

Total phosphorus concentrations in the open waters of the Laurentian Great Lakes are currently at or beneath target concentrations set by international agreement. Despite the success of phosphorus loading controls in remediating nearshore eutrophication problems in the past, nuisance growth of Cladophora has recently returned to the lower Great Lakes. This thesis examines soluble reactive phosphorus (SRP) in a northwestern segment of Lake Ontario to assess whether allochthonous or autochthonous sources of phosphate lead to localized areas of PO43- enrichment that may help to explain the seemingly paradoxical resurgence of Cladophora. As SRP is often an overestimate of PO43- in P-limited waters, measures of SRP made with the standard method were compared with measures of SRP made with modified methods (i.e., using dialysis and magnesium-induced co-precipitation) designed to more accurately measure phosphate when it was expected to be at low concentrations. Measures of SRP made with standard and modified methods did not differ, however, SRP was 1 to 3 orders of magnitude higher than a more sensitive steady-state radiobioassay for PO43- used for comparison in offshore waters. Although the utility of SRP is limited when phosphate concentrations are very low, SRP is useful to measure localized areas of phosphate enrichment, and its relative concentrations can be compared in time and space. To quantify the degree to which allochthonous inputs and dreissenids contribute to PO43- concentrations that permit Cladophora growth, intensive sampling for SRP was carried out prior to, during and following the Cladophora growing season. SRP was higher in the nearshore than offshore and near the mouth of a large tributary and a treated wastewater outfall than in samples from other locations along the shoreline, but only in the spring and autumn. Phosphate turnover times indicated lower P-limitation in the nearshore and near local inputs versus the offshore. Higher concentrations of SRP were measured in samples taken 15 cm and 50 cm above dreissenid mussel-beds than in those obtained at corresponding depths over other substrata and from higher up in the water column through the Cladophora growing season, while Chl a concentrations displayed the reverse trend. These results suggest that PO43- excreted by dreissenids could be more important in time and space than external inputs in supporting nuisance Cladophora growth in the current nearshore environment. Continued research and monitoring of P dynamics in the nearshore combined with model approaches should better predict whether more stringent P controls would be effective in managing Cladophora growth.