Spatial and temporal distribution of gerrid (Heteroptera) and predation on microcrustaceans from a tropical shallow lake / Distribuição espacial e temporal de gerrídeo (Heteroptera) e a predação sobre microcrustáceos de um lago tropical raso

Abstract Predator-prey interactions involving an aquatic insect and zooplanktonic prey of different sizes were investigated to quantify prey mortality exposed to predators. Laboratory experiments were undertaken with the young and adult gerrid Rheumatobates crassifemur to test predation and size selectivity on the cladocerans Daphnia gessneri, Ceriodaphnia richardi, and Bosmina tubicen. Population fluctuations and spatial distribution of the gerrid were also evaluated in a small and shallow Brazilian lake throughout 12 months in fortnightly samples. The insects were more abundant in the littoral (mean density 7.0 ± 1.2 ind.m-2) compared to the limnetic zone. The period with the highest densities was late January to June, in both zones. Predation by young instars on Daphnia and Ceriodaphnia was significant (mean ingestion rate of 1.3 ± 0.1 D. gessneri and 0.7 ± 0.1 C. richardi per predator per hour). Adult insect fed only the large-sized prey (mean ingestion rate of 1.0 ± 0.1 D. gessneri per predator per hour). Young gerrids have greater potential to prey on cladocerans than adults, and size selectivity occurred for both predators. Preference of adults by the larger prey is probably related to difficulties in manipulating smaller planktonic prey, such as Ceriodaphnia. Due to higher densities of insects in the littoral, higher predation on zooplankton in this zone is expected. This study does contribute to a better understanding of trophic interactions in tropical shallow lakes and is the first to investigate predation of a gerrid on cladocerans in laboratory experiments.

Resumo Foram investigadas as interações predador-presa, envolvendo inseto aquático e presas zooplanctônicas de diferentes tamanhos para quantificar a mortalidade das presas expostas a predadores. Experimentos de laboratório foram realizados com jovens e adultos do gerrídeo Rheumatobates crassifemur para testar a predação e a seletividade por tamanho das espécies de cladóceros Daphnia gessneri, Ceriodaphnia richardi e Bosmina tubicen. Flutuações populacionais e a distribuição espacial do gerrídeo também foram avaliadas em um lago brasileiro pequeno e raso durante 12 meses em amostras quinzenais. Os insetos foram mais abundantes no litoral (densidade média 7,0 ± 1,2 ind.m-2) em comparação com a zona limnética. Maiores densidades ocorreram de fim de janeiro a junho, em ambas as zonas. A predação dos jovens foi significativa sobre Daphnia e Ceriodaphnia (taxa de ingestão média 1,3 ± 0,1 D. gessneri and 0,7 ± 0,1 C. richardi por predador por hora). Adultos predaram somente a presa maior (taxa de ingestão média 1,0 ± 0,1 D. gessneri por predador por hora). Gerrídeos jovens têm um maior potencial para predar cladóceros que os adultos, e seletividade por tamanho ocorreu para ambos. A preferência dos adultos somente pela presa maior provavelmente está relacionada à dificuldade em manipular pequenas presas planctônicas como Ceriodaphnia. Devido às maiores densidades de insetos no litoral, é esperada maior predação sobre o zooplâncton nesta zona. Este estudo contribui para uma melhor compreensão sobre interações tróficas em lagos tropicais rasos e é o primeiro a investigar a predação de um gerrídeo sobre cladóceros em experimentos de laboratório.

Distribution and fluctuations of backswimmers (Notonectidae) in a tropical shallow lake and predation on microcrustaceans / Distribuição e flutuação de notonectídeos (Notonectidae) em um lago tropical raso e a predação sobre microcrustáceos

Abstract Notonectids are widely distributed in freshwaters and can prey on zooplankton in temperate lakes. However, its role in structuring the zooplankton community is unknown in tropical lakes. Thus, our objective was to study the notonectid Martarega uruguayensis in a Brazilian tropical shallow lake to evaluate its potential as a zooplankton predator. Its horizontal distribution was analized in the lake throughout one year in fortnightly samplings. Backswimmers were more abundant (mean density 162.9 ± 25.8 ind.m–2) in the cool-dry season, with a strong preference by the littoral zone (mean density 139.9 ± 17.5 ind.m–2). Laboratory experiments were undertaken with young and adult notonectid and the two most abundant cladocerans, Daphnia gessneri and Ceriodaphnia richardi, as prey. Predation by backswimmers in the laboratory showed that only juveniles fed on microcrustaceans (mean ingestion rate of 1.2 ± 0.2 Daphnia and 1.0 ± 0.2 Ceriodaphnia per predator per hour), without size selectivity. Adult insects probably have difficulties in detecting and manipulating small planktonic organisms. On the other hand, young instars might influence zooplankton community, especially in the littoral zone of the lake. This study does contribute to a better understanding of trophic interactions in tropical shallow lakes and is the first to investigate the predation of a notonectid on microcrustaceans from Lake Monte Alegre.

Resumo Notonectídeos são amplamente distribuídos em águas continentais e podem predar zooplâncton em lagos temperados. Porém, seu papel na estruturação de comunidades zooplanctônicas é desconhecido em lagos tropicais. Então, nosso objetivo foi estudar o notonectídeo Martarega uruguayensis em um lago tropical raso brasileiro para avaliar seu potencial como predador do zooplâncton. Sua distribuição horizontal foi analisada no lago durante um ano em coletas quinzenais. Os notonectídeos foram mais abundantes (densidade média 162,9 ± 25,8 ind.m–2) na estação fria-seca, com nítida preferência pela zona litorânea (densidade média 139,9 ± 17,5 ind.m–2). Foram feitos experimentos de laboratório com jovens e adultos de notonectídeo e as duas espécies de cladóceros mais abundantes, Daphnia gessneri and Ceriodaphnia richardi, como presas. No laboratório, somente os jovens predaram os microcrustáceos (taxa média de ingestão 1,2 ± 0,2 Daphnia e 1,0 ± 0,2 Ceriodaphnia por predador por hora), sem haver seletividade. Os insetos adultos provavelmente têm dificuldade em detectar e manipular pequenos organismos planctônicos. Por outro lado, os estádios jovens podem influenciar a comunidade zooplanctônica, especialmente na zona litorânea do lago. Esse estudo contribui para uma melhor compreensão sobre interações tróficas em lagos tropicais rasos e é o primeiro a investigar a predação de um notonectídeo sobre microcrustáceos do Lago Monte Alegre.

Spatial and temporal distribution of gerrid (Heteroptera) and predation on microcrustaceans from a tropical shallow lake.

Predator-prey interactions involving an aquatic insect and zooplanktonic prey of different sizes were investigated to quantify prey mortality exposed to predators. Laboratory experiments were undertaken with the young and adult gerrid Rheumatobates crassifemur to test predation and size selectivity on the cladocerans Daphnia gessneri, Ceriodaphnia richardi, and Bosmina tubicen. Population fluctuations and spatial distribution of the gerrid were also evaluated in a small and shallow Brazilian lake throughout 12 months in fortnightly samples. The insects were more abundant in the littoral (mean density 7.0 ± 1.2 ind.m-2) compared to the limnetic zone. The period with the highest densities was late January to June, in both zones. Predation by young instars on Daphnia and Ceriodaphnia was significant (mean ingestion rate of 1.3 ± 0.1 D. gessneri and 0.7 ± 0.1 C. richardi per predator per hour). Adult insect fed only the large-sized prey (mean ingestion rate of 1.0 ± 0.1 D. gessneri per predator per hour). Young gerrids have greater potential to prey on cladocerans than adults, and size selectivity occurred for both predators. Preference of adults by the larger prey is probably related to difficulties in manipulating smaller planktonic prey, such as Ceriodaphnia. Due to higher densities of insects in the littoral, higher predation on zooplankton in this zone is expected. This study does contribute to a better understanding of trophic interactions in tropical shallow lakes and is the first to investigate predation of a gerrid on cladocerans in laboratory experiments.

Distribution and fluctuations of backswimmers (Notonectidae) in a tropical shallow lake and predation on microcrustaceans.

Notonectids are widely distributed in freshwaters and can prey on zooplankton in temperate lakes. However, its role in structuring the zooplankton community is unknown in tropical lakes. Thus, our objective was to study the notonectid Martarega uruguayensis in a Brazilian tropical shallow lake to evaluate its potential as a zooplankton predator. Its horizontal distribution was analized in the lake throughout one year in fortnightly samplings. Backswimmers were more abundant (mean density 162.9 ± 25.8 ind.m-2) in the cool-dry season, with a strong preference by the littoral zone (mean density 139.9 ± 17.5 ind.m-2). Laboratory experiments were undertaken with young and adult notonectid and the two most abundant cladocerans, Daphnia gessneri and Ceriodaphnia richardi, as prey. Predation by backswimmers in the laboratory showed that only juveniles fed on microcrustaceans (mean ingestion rate of 1.2 ± 0.2 Daphnia and 1.0 ± 0.2 Ceriodaphnia per predator per hour), without size selectivity. Adult insects probably have difficulties in detecting and manipulating small planktonic organisms. On the other hand, young instars might influence zooplankton community, especially in the littoral zone of the lake. This study does contribute to a better understanding of trophic interactions in tropical shallow lakes and is the first to investigate the predation of a notonectid on microcrustaceans from Lake Monte Alegre.

Diel horizontal distribution of microcrustaceans and predators throughout a year in a shallow neotropical lake.

The focus of this study is to investigate if microcrustaceans undergo diel horizontal migration (DHM) in a tropical shallow lake on a yearly basis and analyse the adaptive value regarding predation. Abundance of invertebrate predators, chaoborid larvae and water mites, and microcrustaceans (cladocerans and copepods) were evaluated on a monthly basis in three stations located on a transect during the day and at night. Both invertebrate predators were predominantly pelagic. Cladocerans did not undergo significant DHM, distributing indistinctly onshore and offshore or being mostly pelagic. Nauplii, copepodites of two copepod species and adults of Tropocyclops prasinus meridionalis Kiefer were mostly distributed offshore, and did not perform DHM. The limnological features (temperature, dissolved oxygen, pH, and conductivity) were suitable for the organisms in both zones of the lake. Algal food concentration was a little lower in the littoral than in the limnetic zone during the day, but it seems to be suitable for the organisms. However, as the algae quality was not evaluated, it is not possible to be conclusive concerning its influence. The results indicated that DHM was not performed by the microcrustaceans and is not, therefore, a strategy for decreasing predation by both invertebrates, Chaoborus brasiliensis Theobald and Krendowskia sp., on a yearly basis in this shallow lake.

Experiments on the influence of Chaoborus brasiliensis Theobald, 1901 (Diptera: Chaoboridae) on the diel vertical migration of microcrustaceans from Lake Monte Alegre, Brazil.

The aim of this work was to evaluate whether diel vertical migration of microcrustaceans from Lake Monte Alegre is related to invertebrate predators, namely larvae of Chaoborus brasiliensis. The hypothesis tested in the experiments was that the migratory behaviour of prey would depend on physical contact with Chaoborus brasiliensis larvae or with kairomones released by them. Two experiments were undertaken in the laboratory, with two treatments and three replicates each. Experiments were carried out in acrylic cylinders 1.5 m high and 0.20 m in diameter. Treatments in Experiment I were: A) predator and kairomones present and B) predator and kairomones present, but predator isolated in net tube inside the cylinder, having no physical contact with prey. Treatments in Experiment II were: A) predator and kairomones absent and B) the same as in experiment I. Physical and chemical factors (temperature, dissolved oxygen, pH, electrical conductivity) and food (algae) did not influence vertical distribution of prey, as they were almost homogeneously distributed in the water column in both experiments; food concentrations were not limiting. Adult copepods (Tropocyclops prasinus meridionalis and Thermocyclops decipiens) showed reverse migration in the treatment B of both experiments; the cladoceran Daphnia gessneri migrated nocturnally in the treatment A of the Experiment I; vertical migration was undetectable for copepodites, as well as for Chaoborus larvae. The experiments indicated that Daphnia responded to physical contact with Chaoborus larvae but not to kairomones alone. Adult copepods seemed to be more sensitive to kairomones.

Driving forces of the diel distribution of phytoplankton functional groups in a shallow tropical lake (Lake Monte Alegre, Southeast Brazil).

Phytoplankton vertical and diel dynamics in a small shallow lake (Lake Monte Alegre, Ribeirão Preto, state of São Paulo) were investigated in two climatological periods: July 2001 (cool-dry season) and March 2002 (warm-rainy season). Monte Alegre is a eutrophic reservoir, with a warm polymictic discontinuous circulation pattern. The lake was thermally stratified in both periods, although dissolved oxygen varied less in the cool-dry period. Phytoplankton biomass was higher in the warm-rainy season and the vertical distribution was stratified in both seasons. Flagellate groups (L(m), Y, W(1) and W(2)) and functional groups typical of shallow eutrophic environments (J, X(1) and S(n)) were important throughout the study period. The lake's thermal pattern strongly influenced the vertical distribution of the phytoplankton community in both periods. Biomass, functional groups and size classes of phytoplankton also were determined by the presence of more efficient herbivores in the lake, especially during the cool-dry period when phytoplankton biomass decreased.

Driving forces of the diel distribution of phytoplankton functional groups in a shallow tropical lake (Lake Monte Alegre, Southeast Brazil) / Fatores direcionadores da distribuição nictemeral de grupos funcionais fitoplanctônicos de um lago brasileiro raso e tropical

Phytoplankton vertical and diel dynamics in a small shallow lake (Lake Monte Alegre, Ribeirão Preto, state of São Paulo) were investigated in two climatological periods: July 2001 (cool-dry season) and March 2002 (warm-rainy season). Monte Alegre is a eutrophic reservoir, with a warm polymictic discontinuous circulation pattern. The lake was thermally stratified in both periods, although dissolved oxygen varied less in the cool-dry period. Phytoplankton biomass was higher in the warm-rainy season and the vertical distribution was stratified in both seasons. Flagellate groups (Lm, Y, W1 and W2) and functional groups typical of shallow eutrophic environments (J, X1 and Sn) were important throughout the study period. The lake's thermal pattern strongly influenced the vertical distribution of the phytoplankton community in both periods. Biomass, functional groups and size classes of phytoplankton also were determined by the presence of more efficient herbivores in the lake, especially during the cool-dry period when phytoplankton biomass decreased.

As dinâmicas vertical e nictemeral do fitoplâncton de um lago pequeno e raso (Lago Monte Alegre, Ribeirão Preto, SP) foram investigadas em dois períodos climatológicos: julho/2001 (estação fria-seca) e março/2002 (estação quente-chuvosa). O lago esteve estratificado termicamente nos dois períodos de estudo, porém menores variações do oxigênio dissolvido foram observadas no período frio-seco. Maiores biomassas fitoplanctônicas foram registradas na estação quente-chuvosa e a distribuição vertical esteve estratificada nos dois períodos climatológicos. Grupos de flagelados (Lm, Y, W1 e W2), juntamente com grupos funcionais típicos de ambientes rasos e eutróficos (J, X1 e Sn), foram importantes em todo o estudo. O padrão térmico do lago teve influência na distribuição vertical da comunidade fitoplanctônica nos períodos estudados. Biomassa, grupos funcionais e classes de tamanho do fitoplâncton também foram influenciados pela presença de herbívoros mais eficientes, principalmente durante o período frio-seco, quando ocorreram menores biomassas do fitoplâncton.

Influence of seston quantity and quality on growth of tropical cladocerans.

The seston of the small, shallow, and tropical lake Monte Alegre was tested for quantity and quality for cladocerans by growth bioassays, which were carried out in spring (Daphnia gessneri and Moina micrura), summer (D. gessneri, M. micrura, Ceriodaphnia cornuta, and Simocephalus mixtus), and winter (D. gessneri and D. ambigua). Cohorts of newborns originating from ovigerous females collected in the lake or from laboratory cultures were submitted, at a room temperature of 23 degrees C to the following treatments: (1) the chlorophytes Ankistrodesmus falcatus and/or Scenedesmus spinosus; (2) lake seston; and (3) lake seston + chlorophytes. Growth rate, clutch size, and fecundity were evaluated. Seston alone was not the best food for promoting cladoceran growth. There were seasonal differences in food quantity and quality with spring and summer seston being better for growth than that of the winter. Adding chlorophytes to the seston increased clutch size and fecundity for most species in summer and winter, but not in spring. Energy limitation seems to be the most important factor influencing cladoceran growth in summer and especially in winter.

Influence of seston quantity and quality on growth of tropical cladocerans

. (O séston do lago Monte Alegre, um lago tropical pequeno e raso, foi testado em relação à quantidade e qualidade para os cladóceros por meio de experimentos de crescimento realizados na primavera (Daphnia gessneri e Moina micrura), no verão (D. gessneri, M. micrura, Ceriodaphnia cornuta e Simocephalus mixtus) e no inverno (D. gessneri e D. ambigua). Coortes de recém-nascidos oriundos de fêmeas ovígeras coletadas no lago ou de culturas de laboratório foram submetidas à temperatura de 23ºC aos seguintes tratamentos: (1) as clorofíceas Ankistrodesmus falcatus e/ou Scenedesmus spinosus, (2) séston do lago e (3) séston + clorofíceas. Foram avaliados a taxa de crescimento, o tamanho da ninhada e a fecundidade. O séston sozinho não foi o melhor alimento para promover o crescimento dos cladóceros. Houve diferenças sazonais quanto à quantidade e à qualidade do alimento, sendo o séston da primavera e o do verão melhores para promover o crescimento que o do inverno. A adição de clorofíceas ao séston aumentou o tamanho da ninhada e a fecundidade para a maioria das espécies no verão e no inverno, mas não na primavera. A limitação de energia parece ser o fator mais importante para o crescimento de cladóceros no verão e, especialmente, no inverno.

Diel vertical migration of copepods in a Brazilian lake: a mechanism for decreasing risk of Chaoborus predation?

A comparison between two studies on diel vertical migration of two cyclopoid copepod species, in Lake Monte Alegre, undertaken in 1985/86 and 1999, revealed a change in their migratory behavior. In summer, during a period of marked stratification with low dissolved oxygen near the bottom, the organisms avoided the deepest layers, and migration was nocturnal or undetectable, in both periods. On other occasions, with partial or total circulation in the lake, a weak twilight migration of copepodites and adults in 1985 was replaced by the reverse in 1999. Differences were found among stages, with the weakest or undetectable migration being observed for nauplii. The migratory pattern change for copepodites and adults might be related with a recent predation pressure increase by Chaoborus larvae on copepods, after the virtual disappearance of their main cladoceran prey.

Diel vertical migration of copepods in a Brazilian lake: a mechanism for decreasing risk of Chaoborus predation?

A comparação entre dois estudos sobre migração vertical diária de duas espécies de copépodos ciclopóides no Lago Monte Alegre, realizados entre 1985/86 e 1999, revelou mudança em seu comportamento migratório. No verão, durante período de marcada estratificação, com pouco oxigênio dissolvido próximo ao fundo, as camadas mais profundas foram evitadas pelos organismos e a migração foi noturna ou não ocorreu. Em outras ocasiäes, com circulação parcial ou total do lago, uma fraca migração crepuscular de copepoditos e adultos, em 1985, foi substituída por reversa em 1999. Foram encontradas diferenças entre os estágios, sendo que os náuplios não migraram ou migraram fracamente. A mudança do padrão migratório de copepoditos e adultos poderia estar relacionada a recente aumento da predação de copépodos por larvas de Chaoborus, depois do desaparecimento virtual do cladócero, sua presa principal.

Water mite predation on zooplankton of a tropical lake

Water mite (Krendowskia sp.) predation on zooplanktonic species from Lake Monte Alegre (Säo Paulo State, Brazil) was tested in laboratory experiments. One cladoceran, Daphnia gessneri, two copepods, Tropocyclops prasinus and Thermocyclops decipiens, and Chaoborus larvae III and IV were supplied as prey to adult mites. The cladoceran Ceriodaphnia cornuta was used in the experiment with mite nymphs. Only D.gessneri was preyed on by mites, at an average rate of 7 ind./mite/day, at 23-24ºC. Although Chaoborus was not eaten by the predator in the experiment, in direct observations older larvae were seen to be seized by several mites, both predators and prey sinking to the bottom of the vessel. Adult Daphnia was grabbed by its dorsal side, swimming together with the mite. There was an increase in predatory activity two hours after sunset

Early life stages of fish and the relationships with zooplankton in a tropical Brazilian reservoir: Lake Monte Alegre.

For evaluating the trophic relationship between early life stages of fish and zooplankton in Lake Monte Alegre, fish distribution and feeding habits have been studied in areas with and without macrophytes. In the first of these areas, 356 specimens belonging to 8 species, mostly juveniles, were caught by a sieve. Another 35 specimens, belonging to 4 species, were caught by gill nets and seine in areas lacking macrophytes. Their diets were composed of aquatic insects, microcrustaceans, rotifers, detritus, and other aquatic invertebrates. Microcrustacean prey were mainly littoral or benthic dwellers, found in the highest proportion in specimens of 7-20 mm SL. Most fish species have parental care, which could explain the absence of planktonic larvae. Early life stages of fish do not exert a predation pressure on the lake zooplankton.

Early life stages of fish and the relationships with zooplankton in a tropical Brazilian reservoir: Lake Monte Alegre

For evaluating the trophic relationship between early life stages of fish and zooplankton in Lake Monte Alegre, fish distribution and feeding habits have been studied in areas with and without macrophytes. In the first of these areas, 356 specimens belonging to 8 species, mostly juveniles, were caught by a sieve. Another 35 specimens, belonging to 4 species, were caught by gill nets and seine in areas lacking macrophytes. Their diets were composed of aquatic insects, microcrustaceans, rotifers, detritus, and other aquatic invertebrates. Microcrustacean prey were mainly littoral or benthic dwellers, found in the highest proportion in specimens of 7-20 mm SL. Most fish species have parental care, which could explain the absence of planktonic larvae. Early life stages of fish do not exert a predation pressure on the lake zooplankton

Water mite predation on zooplankton of a tropical lake.

Water mite (Krendowskia sp.) predation on zooplanktonic species from Lake Monte Alegre (São Paulo State, Brazil) was tested in laboratory experiments. One cladoceran, Daphnia gessneri, two copepods, Tropocyclops prasinus and Thermocyclops decipiens, and Chaoborus larvae III and IV were supplied as prey to adult mites. The cladoceran Ceriodaphnia cornuta was used in the experiment with mite nymphs. Only D. gessneri was preyed on by mites, at an average rate of 7 ind./mite/day, at 23-24 degrees C. Although Chaoborus was not eaten by the predator in the experiment, in direct observations older larvae were seen to be seized by several mites, both predators and prey sinking to the bottom of the vessel. Adult Daphnia was grabbed by its dorsal side, swimming together with the mite. There was an increase in predatory activity two hours after sunset.

Feeding habits of Chaoboridae larvae in a tropical Brazilian reservoir

The diet of the four larval instars of a Chaoboridae species of Lake Monte Alegre, was evaluated through the analysis of the crop content of individuals caught at night in the lake. Peridinium and zooflagellates were the major components of the diet of instars I and II, whereas microcrustaceans, especially the cladoceran Bosmina, were most important for late instars. When the preferential prey decreased, in the course of the year, the diet was more diversified, increasing the ingestion of Peridinium by instars III and IV. There is a relationship between the mouth diameter of the larvae and the maximum width, height, or diameter of the ingested organisms

Feeding habits of Chaoboridae larvae in a tropical Brazilian reservoir.

The diet of the four larval instars of a Chaoboridae species of Lake Monte Alegre, was evaluated through the analysis of the crop content of individuals caught at night in the lake. Peridinium and zooflagellates were the major components of the diet of instars I and II, whereas microcrustaceans, especially the cladoceran Bosmina, were most important for late instars. When the preferential prey decreased, in the course of the year, the diet was more diversified, increasing the ingestion of Peridinium by instars III and IV. There is a relationship between the mouth diameter of the larvae and the maximum width, height, or diameter of the ingested organisms.

Production of the rotifer Brachionus plicatilis (Ploimida: Brachionidae) in a Brazilian coastal lagoon.

Secondary production, density and biomass fluctuations, and some reproductive aspects of the rotifer Brachionus plicatilis were studied weekly for two years in a Brazilian coastal lagoon. Food (and secondarily temperature) seemed to be the main factor involved in the summer increments. In the first summer, a mass mortality of fish caused changes in the phytoplankton, which switched from the dominance of Cyanophyceae to Chlorophyceae and other groups. For two months, production, density and biomass were relatively high until the recovery of the Cyanophyceae dominance. In the second summer, peaks of shorter duration and higher values of production, in comparison to the first one, were found. The contribution of resting eggs hatching, in the first summer, and the increment in parthenogenetic reproduction, in the second summer, would account for the population increases observed. Males and mictic females occurred when densities were high.