Methods for selection of Daphnia resting eggs: the influence of manual decapsulation and sodium hypoclorite solution on hatching rates / Técnicas para seleção de ovos de resistência de Daphnia: influência do desencapsulamento manual e imersão em solução de hipoclorito nas taxas de eclosão

Abstract Cladocerans are able to produce resting eggs inside a protective resistant capsule, the ephippium, that difficults the visualization of the resting eggs, because of the dark pigmentation. Therefore, before hatching experiments, methods to verify viable resting eggs in ephippia must be considered. This study aimed to evaluate the number of eggs per ephippium of Daphnia from two tropical aquatic ecosystems and the efficiency of some methods for decapsulating resting eggs. To evaluate the influence of methods on hatching rates, three different conditions were tested: immersion in sodium hypochlorite, manually decapsulated resting eggs and intact ephippia. The immersion in hypochlorite solution could evaluate differences in numbers of resting eggs per ephippium between the ecosystems studied. The exposure to sodium hypochlorite at a concentration of 2% for 20 minutes was the most efficient method for visual evaluation and isolation of the resting eggs. Hatching rate experiments with resting eggs not isolated from ephippia were underestimated (11.1 ± 5.0%), showing the need of methods to quantify and isolate viable eggs. There were no differences between the hatching rate of resting eggs submitted to hypochlorite solution (47.2 ± 7.34%) and manually decapsulated (53.7 ± 13.24%). However, the immersion in hypochlorite was a more efficient technique, faster and not requiring manual ability.

Resumo Cladóceros são capazes de produzir ovos de resistência envoltos por uma cápsula protetora, o efípio, que dificulta a visualização do número de ovos, devido à pigmentação escura. Assim, antes da condução de experimentos de eclosão, métodos para determinar o número de ovos de resistência viáveis no interior do efípio devem ser utilizados. Este estudo teve como objetivo avaliar o número de ovos por efípio de Daphnia de dois ecossistemas aquáticos tropicais e a eficiência de alguns métodos de desencapsulamento de ovos de resistência. Para avaliar a influência dos métodos na taxa de eclosão foram testadas três condições: imersão em hipoclorito de sódio, ovos de resistência desencapsulados manualmente e efípios intactos. A imersão em solução de hipoclorito permitiu avaliar as diferenças no número de ovos de resistência por efípios entre os ecossistemas. A exposição ao hipoclorito de sódio a uma concentração de 2% durante 20 minutos foi o método mais eficiente para a avaliação visual e isolamento dos ovos de resistência. A taxa de eclosão de ovos não isolados de efípios foi subestimada (11,1 ± 5,0%), mostrando a necessidade de métodos para quantificação e isolamento dos ovos. Não foi verificada diferença de eclosão entre os ovos isolados a partir da imersão em solução de hipoclorito (47,2 ± 7,34%) e manualmente desencapsulados (53,7 ± 13,24%). Entretanto, a imersão em hipoclorito é uma técnica mais eficiente, pois além de mais rápida, não necessita de destreza manual.

Methods for selection of Daphnia resting eggs: the influence of manual decapsulation and sodium hypoclorite solution on hatching rates.

Cladocerans are able to produce resting eggs inside a protective resistant capsule, the ephippium, that difficults the visualization of the resting eggs, because of the dark pigmentation. Therefore, before hatching experiments, methods to verify viable resting eggs in ephippia must be considered. This study aimed to evaluate the number of eggs per ephippium of Daphnia from two tropical aquatic ecosystems and the efficiency of some methods for decapsulating resting eggs. To evaluate the influence of methods on hatching rates, three different conditions were tested: immersion in sodium hypochlorite, manually decapsulated resting eggs and intact ephippia. The immersion in hypochlorite solution could evaluate differences in numbers of resting eggs per ephippium between the ecosystems studied. The exposure to sodium hypochlorite at a concentration of 2% for 20 minutes was the most efficient method for visual evaluation and isolation of the resting eggs. Hatching rate experiments with resting eggs not isolated from ephippia were underestimated (11.1 ± 5.0%), showing the need of methods to quantify and isolate viable eggs. There were no differences between the hatching rate of resting eggs submitted to hypochlorite solution (47.2 ± 7.34%) and manually decapsulated (53.7 ± 13.24%). However, the immersion in hypochlorite was a more efficient technique, faster and not requiring manual ability.

Loss of biodiversity in a conservation unit of the Brazilian Atlantic Forest: the effect of introducing non-native fish species.

The introduction of species has become an important problem for biodiversity and natural ecosystem conservation. The lake system of the middle Rio Doce (MG, Brazil) comprises c. 200 lakes at various conservation states, of which 50 are located within the Rio Doce State Park (PERD). Previous studies had verified several of these lakes suffered non-native fishes introductions and the presence of these species needs for the implementation of actions aiming at not only their control but also the preservation of the native species. This study discusses the effects of non-native fish species in the largest conservation unit of Atlantic Forest in Minas Gerais, southeast of Brazil, using data from 1983 to 2010 distributed as follow: data prior to 2006 were obtained from previous studies, and data from September 2006 to July 2010 were obtained in Lake Carioca at four sampling stations using gillnets, seine nets and sieve. A total of 17 fish species was collected (2006-2010) of which five were introduced species. Among the small to medium size native species (30 to 2000 mm standard length) seven had disappeared, two are new records and one was recaptured. The non-native species Cichla kelberi (peacock bass) and Pygocentrus nattereri (red piranha) are within the most abundant captured species. Integrated with other actions, such as those preventing new introductions, a selective fishing schedule is proposed as an alternative approach to improve the conservation management actions and the local and regional biodiversity maintenance.

Loss of biodiversity in a conservation unit of the Brazilian Atlantic Forest: the effect of introducing non-native fish species / Perda de biodiversidade em uma unidade de conservação da Mata Atlântica brasileira: efeitos da introdução de espécies não nativas de peixes

Abstract The introduction of species has become an important problem for biodiversity and natural ecosystem conservation. The lake system of the middle Rio Doce (MG, Brazil) comprises c. 200 lakes at various conservation states, of which 50 are located within the Rio Doce State Park (PERD). Previous studies had verified several of these lakes suffered non-native fishes introductions and the presence of these species needs for the implementation of actions aiming at not only their control but also the preservation of the native species. This study discusses the effects of non-native fish species in the largest conservation unit of Atlantic Forest in Minas Gerais, southeast of Brazil, using data from 1983 to 2010 distributed as follow: data prior to 2006 were obtained from previous studies, and data from September 2006 to July 2010 were obtained in Lake Carioca at four sampling stations using gillnets, seine nets and sieve. A total of 17 fish species was collected (2006-2010) of which five were introduced species. Among the small to medium size native species (30 to 2000 mm standard length) seven had disappeared, two are new records and one was recaptured. The non-native species Cichla kelberi (peacock bass) and Pygocentrus nattereri (red piranha) are within the most abundant captured species. Integrated with other actions, such as those preventing new introductions, a selective fishing schedule is proposed as an alternative approach to improve the conservation management actions and the local and regional biodiversity maintenance.

Resumo A introdução de espécies tornou-se um importante problema para a conservação da biodiversidade e dos ecossistemas naturais. O sistema de lagos do médio Rio Doce (MG, Brasil) compreende cerca de 200 lagos em vários estados de conservação, dos quais 50 estão localizados dentro do Parque Estadual do Rio Doce (PERD). Estudos anteriores demonstraram que vários desses lagos sofreram introduções de peixes não nativos. Este estudo discute os efeitos das espécies de peixes não nativos na maior unidade de conservação da Mata Atlântica em Minas Gerais, sudeste do Brasil, utilizando dados de 1983 até 2010, distribuídos da seguinte forma: dados de 1983 a 2005 obtidos de estudos realizados anteriormente, e dados de setembro de 2006 a julho de 2010 obtidos no lago Carioca, utilizando redes de emalhar, redes de arrasto e peneira. Um total de 17 espécies de peixes foi coletado (2006-2010), sendo cinco introduzidas. Dentre estas, Cichla kelberi (tucunaré) e Pygocentrus nattereri (piranha vermelha) estiveram dentre as mais abundantes. Entre as espécies nativas de pequeno a médio porte (30-2000 mm de comprimento padrão), duas foram novos registros e uma foi recapturada. O desaparecimento de sete destas espécies, registradas em estudos anteriores, e a baixa representatividade nas amostragens da maioria das espécies remanescentes, evidencia o alto impacto da invasão de peixes na comunidade estudada. Considerando a importância de ações de manejo de espécies não nativas visando à conservação e manutenção da biodiversidade local e regional, propõem-se a realização de pescas seletivas intensivas, juntamente com ações que previnam novas introduções, incluindo ações educativas junto às comunidades locais.

Cyanobacteria bloom: selective filter for zooplankton?

The Ibirité reservoir is an urban and eutrophic environment, with regular occurrences of cyanobacteria blooms. The reservoir is warm monomict and remains stratified most of the year, circulating in the dry season (winter). During the hydrological cycle of October/07 to October/08 there were four scenarios with different environmental conditions, which influenced the structure of the zooplankton community, as confirmed in a previous study. Changes in the zooplankton community structure between the scenarios were studied, aiming at analyzing the stability and persistence of this community. The Spearman's coefficient of correlation was used to measure the stability; the persistence was evaluated through a cluster analysis and changes in community composition were estimated by the "temporal" ß diversity index. Considering the distribution patterns of abundance, the community was stable only in the transition between scenarios 1 and 2 (n = 30, r = 0.71, p = 0.00001), when there were no cyanobacteria blooms. The persistence of zooplankton between the scenarios was low, showing a distinct species composition for each scenario. The highest variations in species composition, observed by the values of temporal ß diversity index, were the transitions between scenarios 3-0 (1.45) and 0-1 (1.05), and the lowest variations occurred in the transition between scenarios 1-2 (0.57). The results suggest that the cyanobacteria blooms at Ibirité reservoir are be acting as "selective filters", and are, thus, disturbances with sufficient ability to change the structure of the zooplankton community.

Seasonal dynamics of Daphnia laevis Birge, 1878 ephippia in a tropical lake with a description of a new methodology for in situ evaluation.

The effect of dormancy in zooplankton populations is still unknown, largely because of the lack of methods to estimate hatching and production of the dormant stages. This study aimed to compare the production and hatching rates of ephippia of Daphnia laevis between thermal stratification and mixing periods in Jacaré Lake (Middle Rio Doce, Minas Gerais, Brazil). For this, we collected ephippia on the sediment with core sampler and we created a device called the "Ephippial Collector". There was a significant difference in ephippia hatching in situ between stratification and mixing periods (Pearson's Chi-squared test p <0.001), being higher in the second one. Significant differences in the hatching rates between periods was observed in the laboratory only for ephippia collected with Ephippial Collectors (Pearson's Chi-squared test p <0.001), being higher during the mixing period (∼ 8%). The core sample allows the collection of a certain fraction of the sediment that may contain a mixture of ephippia produced in different periods, i.e., may contain old and not viable ephippia, which masks the hatching rate. Thus, seasonality in hatching rates of ephippia was reported only by Ephippial Collectors. The higher hatching rate observed during the mixing period in the lake suggests that individuals hatched from ephippia may contribute to the increase in the population of D. laevis in the water column at this time.

Length-weight relationships and biomass of the main microcrustacean species of two large tropical reservoirs in Brazil.

Length-weight equations were determined for the main microcrustacean species of Três Marias and Furnas reservoirs (state of Minas Gerais, Brazil), and evaluated the fluctuations of their biomass. We examined the following species: Thermocyclops minutus, Bosminopsis deitersi, Bosmina hagmanni, Ceriodaphnia cornuta, and Moina minuta (Três Marias Reservoir), and Notodiaptomus henseni, Daphnia ambigua, Ceriodaphnia silvestrii, Diaphanosoma spinulosum, D. fluviatile, and Bosmina freyi (Furnas Reservoir). Dry weight was obtained in a microbalance for each size class (Cladocera) or developmental stage (Copepoda). Microcrustacean mean biomass varied from 5.76 mg DW.m(-3) and 20.36 mg DW.m(-3) (Furnas Reservoir) and from 3.75 mg DW.m(-3) and 18.14 mg DW.m(-3) (Três Marias Reservoir). Significant differences (p < 0.000) between seasons were registered with higher biomass during the rainy seasons. Thermocyclops minutus was the most important species in Três Marias, whereas in Furnas, N. henseni contributed in equal proportion. In Furnas, the higher cladoceran biomass was explained by the contribution of larger-sized species, such as D. ambigua, C. silvestrii, D. spinulosum, and D. fluviatile. Even though both reservoirs are considered oligotrophic, there were significant differences in the estimated microcrustacean biomass, which reinforces the importance of this parameter for the description of the real contribution of each species in the community.

Limnological changes in Dom Helvécio Lake (South-East Brazil): natural and anthropogenic causes.

In order to verify changes in physical, chemical and biological features of Dom Helvécio Lake (South-East Brazil), this study compares previous and recent data obtained from limnological investigations over three decades. Plankton species composition and density together with environmental data from 1978, 1983, and during 1999 to 2006 showed changes driven by seasonal climatic forces or by anthropogenic causes. Changes in diversity are shown as a consequence of alternation of rainy and dry periods and introduction of exotic fish species. Finally, suggestions are made for improving conservation efforts in the area, through planned actions and monitoring programmes.

The inverted trophic cascade in tropical plankton communities: impacts of exotic fish in the Middle Rio Doce lake district, Minas Gerais, Brazil / A cascata trófica invertida em comunidades planctônicas tropicais: impactos da introdução de peixes exóticos no distrito de lagos do médio rio Doce, MG

The present study deals with the ecological impacts of the introduction of two alien species of piscivorous fish in several lakes of the Middle Rio Doce lake district in Minas Gerais, Brazil. It was demonstrated that these effects were not restricted only to the fish community. The introduction of the predatory red piranha Pygocentrus nattereri and the tucunaré Cichla cf. ocellaris caused not only a sharp decrease in the number of native fish species, but also major shifts in other trophic levels. Just after the fish were introduced, most lakes began to show conspicuous changes in phytoplankton species composition, in which Cyanophyceae gradually came to dominate. The zooplankton community lost several species, and in some cases, such as Lake Carioca, all the cladoceran species disappeared. On the other hand, invertebrate predators, represented by the dipteran Chaoboridae, boomed in the lake, with higher densities of exotic species, probably as a result of the "ecological release" by reduction of the original fish fauna. There was a general trend of species loss in different trophic levels. All these changes are apparently associated with decreases in water quality. The present situation in these lakes demands new approaches to the management and conservation of these ecosystems.

O presente estudo trata dos impactos ecológicos da introdução de duas espécies invasoras de peixes piscívoros em diversos lagos da região lacustre do médio rio Doce em Minas Gerais, Brasil. Demonstrou-se que estes efeitos não se restringiram às comunidade de peixes. A introdução dos predadores Pygocentrus nattereri (piranha-vermelha) e Cichla cf. ocelaris (tucunaré) não só causou uma forte redução no número de espécies de peixes nativos, como também mudanças nos níveis tróficos inferiores. Pouco depois das introduções, a maioria dos lagos começou a mostrar alterações na comunidade fitoplanctônica, tais como o aparecimento da dominância de Cyanophyceae. A comunidade zooplanctônica perdeu diversas espécies e, em alguns casos, houve o desaparecimento de todas as espécies de cladóceros limnéticos, como é o caso da lagoa Carioca. Por outro lado, predadores invertebrados, representados pelos dípteros da família Chaoboridae, floresceram nos lagos com maiores densidades de espécies exóticas de peixes, provavelmente como resultado da "liberação ecológica" causada pela redução da ictiofauna original. Além de uma tendência geral de perda de espécies em diferentes níveis tróficos, outras mudanças estão aparentemente associadas com a redução da qualidade de água. Dessa forma, esses ecossistemas estão necessitando urgentemente de novas abordagens nas estratégias de manejo e conservação.

Zooplankton in littoral waters of a tropical lake: a revisited biodiversity / Zooplâncton da região litorânea de um lago tropical: a biodiversidade revisitada

The present study was carried out in Lake Dom Helvécio, in the state of Minas Gerais, with two main objectives: to demonstrate the contribution of the littoral zone, in order to better characterize zooplankton fauna; and to assess the distribution of zooplankton species in different habitats, i.e., the littoral zone with and without aquatic vegetation. The samples were collected in February and July 2006, throughout the littoral zone of the lake, in areas with and without aquatic vegetation. We identified a total of 188 species, of which 130 are new records for Lake Dom Helvécio. One hundred and eighty-four species were identified in the littoral zone with aquatic vegetation, and 117 in the zone with no vegetation. The higher zooplankton richness in areas of the littoral zone with aquatic vegetation can be related to the greater environmental heterogeneity. Compared to previous studies on the littoral zones of lakes along the middle River Doce, the present study expended greater sampling effort, and identified many more species. In relation to biological conservation, this study demonstrated the importance of the littoral zone for better characterization and conservation of the zooplankton fauna, especially when it is colonized by aquatic vegetation. Underestimating the richness of species may provide inaccurate data on the biota, as well as on the ecological conditions in an environment.

O presente estudo foi realizado no lago Dom Helvécio (MG) tendo como principais objetivos: demonstrar a contribuição da região litorânea para uma melhor caracterização da fauna zooplanctônica, avaliar a distribuição destas espécies em diferentes habitats (região litorânea com e sem vegetação aquática). As amostras foram coletadas nos meses de fevereiro e julho de 2006, ao longo da região litorânea, em pontos amostrais com e sem vegetação aquática. Um total de 188 espécies foi identificado. Desses organismos, 130 representaram novos registros para o lago Dom Helvécio. Foram identificadas 184 espécies na região litorânea com vegetação aquática e 117 na região litorânea sem vegetação. A maior riqueza em espécies zooplanctônicas na região litorânea com vegetação aquática está relacionada com a maior heterogeneidade ambiental. Quando comparado aos estudos realizados anteriormente na região litorânea de lagos do médio rio Doce, este estudo destaca-se pelo maior esforço amostral e número de espécies identificadas. No tocante à conservação biológica, o presente estudo mostrou a importância da região litorânea, especialmente quando colonizada por vegetação aquática para uma melhor caracterização e conservação da fauna zooplanctônica. A riqueza em espécies subestimada pode fornecer dados irreais sobre a biota, bem como das condições ecológicas de um ambiente.

The inverted trophic cascade in tropical plankton communities: impacts of exotic fish in the Middle Rio Doce lake district, Minas Gerais, Brazil.

The present study deals with the ecological impacts of the introduction of two alien species of piscivorous fish in several lakes of the Middle Rio Doce lake district in Minas Gerais, Brazil. It was demonstrated that these effects were not restricted only to the fish community. The introduction of the predatory red piranha Pygocentrus nattereri and the tucunaré Cichla cf. ocellaris caused not only a sharp decrease in the number of native fish species, but also major shifts in other trophic levels. Just after the fish were introduced, most lakes began to show conspicuous changes in phytoplankton species composition, in which Cyanophyceae gradually came to dominate. The zooplankton community lost several species, and in some cases, such as Lake Carioca, all the cladoceran species disappeared. On the other hand, invertebrate predators, represented by the dipteran Chaoboridae, boomed in the lake, with higher densities of exotic species, probably as a result of the 'ecological release' by reduction of the original fish fauna. There was a general trend of species loss in different trophic levels. All these changes are apparently associated with decreases in water quality. The present situation in these lakes demands new approaches to the management and conservation of these ecosystems.

Zooplankton in littoral waters of a tropical lake: a revisited biodiversity.

The present study was carried out in Lake Dom Helvécio, in the state of Minas Gerais, with two main objectives: to demonstrate the contribution of the littoral zone, in order to better characterize zooplankton fauna; and to assess the distribution of zooplankton species in different habitats, i.e., the littoral zone with and without aquatic vegetation. The samples were collected in February and July 2006, throughout the littoral zone of the lake, in areas with and without aquatic vegetation. We identified a total of 188 species, of which 130 are new records for Lake Dom Helvécio. One hundred and eighty-four species were identified in the littoral zone with aquatic vegetation, and 117 in the zone with no vegetation. The higher zooplankton richness in areas of the littoral zone with aquatic vegetation can be related to the greater environmental heterogeneity. Compared to previous studies on the littoral zones of lakes along the middle River Doce, the present study expended greater sampling effort, and identified many more species. In relation to biological conservation, this study demonstrated the importance of the littoral zone for better characterization and conservation of the zooplankton fauna, especially when it is colonized by aquatic vegetation. Underestimating the richness of species may provide inaccurate data on the biota, as well as on the ecological conditions in an environment.

On the natural diet of Daphnia laevis in the eutrophic Pampulha reservoir (Belo Horizonte, Minas Gerais)

The aim of this study was to assess the major food items ingested by adult specimens of Daphnia laevis within the eutrophic Pampulha reservoir in Belo Horizonte, Minas Gerais, Brazil. The gut content was analyzed after addition of sodium hypochlorite and also through the examination of dissected guts under scanning electron microscopy. The results showed that Chlorophyceae was the main food item ingested, representing c. 80.5 percent of the total ingested food. Moreover, Eutetramorus fottii, Coelastrum pseudomicroporum and Oocystis lacustris, the dominant phytoplankton species within the reservoir, were the most frequent cells found in the gut contents. Euglenophyta also represented an important food item accounting for 15 percent of the ingested material, including mainly Trachelomonas volvocina and Euglena oxyuris, although less abundant in the reservoir (< 10 percent of total phytoplankton). Blue-green algae occurred at much lower percentages in the guts than in the phytoplankton. A small amount of undigested Microcystis aeruginosa colonies were also found in the gut content of D. laevis. Scanning electron microscopy results showed that, besides phytoplankton cells, a great amount of abiogenic material was also ingested. The amount of this inorganic material increased considerably in the tract (from 15 percent to 75 percent of the gut content), when a peak of D. laevis was observed in the reservoir. Our assumption is that the ingestion of this inorganic material can be a strategy used by D. laevis to obtain additional food supply

On the natural diet of Daphnia laevis in the eutrophic Pampulha reservoir (Belo Horizonte, Minas Gerais).

The aim of this study was to assess the major food items ingested by adult specimens of Daphnia laevis within the eutrophic Pampulha reservoir in Belo Horizonte, Minas Gerais, Brazil. The gut content was analyzed after addition of sodium hypochlorite and also through the examination of dissected guts under scanning electron microscopy. The results showed that Chlorophyceae was the main food item ingested, representing c. 80.5% of the total ingested food. Moreover, Eutetramorus fottii, Coelastrum pseudomicroporum and Oocystis lacustris, the dominant phytoplankton species within the reservoir, were the most frequent cells found in the gut contents. Euglenophyta also represented an important food item accounting for 15% of the ingested material, including mainly Trachelomonas volvocina and Euglena oxyuris, although less abundant in the reservoir (< 10% of total phytoplankton). Blue-green algae occurred at much lower percentages in the guts than in the phytoplankton. A small amount of undigested Microcystis aeruginosa colonies were also found in the gut content of D. laevis. Scanning electron microscopy results showed that, besides phytoplankton cells, a great amount of abiogenic material was also ingested. The amount of this inorganic material increased considerably in the tract (from 15% to 75% of the gut content), when a peak of D. laevis was observed in the reservoir. Our assumption is that the ingestion of this inorganic material can be a strategy used by D. laevis to obtain additional food supply.

Avaliaçäo da comunidade zooplanctônica na regiäo litorânea de dois lagos naturais no Médio Rio Doce / Evaluation of the zooplankton community in the litoral zone of two natural lakes in the middle Rio Doce, Brazil

A avaliaçäo quantitativa do zooplâncton na regiäo litorânea de dois lagos com diferentes usos de suas bacias de drenagem - Mata Atlântica em regeneraçäo (lago Pedra) e reflorestamento com Eucalyptus spp. (lago Hortência), foi investigada em agosto/92 e janeiro/93. No lago Pedra foram identificadas 30 espécies na estaçäo de seca (agosto/92) e 27, na estaçäo chuvosa (janeiro/93) e no lago Hortência, 29 na seca e 28 na estaçäo chuvosa. No lago Pedra, a densidade total de organismos foi de 192,2 org/l na estaçäo de seca e 453,7 org/l na estaçäo de chuvas. No lago Hortência, foi de 624,6 e 484,4 org/l na seca e chuva, respectivamente. Os grupos dominantes no lago Pedra foram Copepoda (57,7 por cento) e Cladocera (21,0 por cento) na seca e Copepoda (32,7 por cento) e Protozoa (57,7 por cento) na chuva. No lago Hortência, Copepoda foi o grupo dominante nos dois períodos amostrados (47,9 por cento na seca e 49,4 por cento na chuva), seguidos por Rotifera (26,0 por cento na seca e 33,8 por cento no período chuvoso). Apesar da semelhança observada na composiçäo da comunidade zooplanctônica entre os dois ambientes a quantidade e qualidade do material alóctone e os efeitos da velocidade de sua decomposiçäo poderiam explicar as diferenças nas densidades dos organismos entre lagos e períodos