Reservoir longitudinal gradient promotes ordered losses on diversity and density of Ephemeroptera community / O gradiente longitudinal de um reservatório promove perdas ordenadas na diversidade e densidade da comunidade de Ephemeroptera

Abstract Reservoir operations alter, eliminate or restrain the natural hydrologic cycles. Biotic community has become subject to these non-cyclic events, responding by reducing the species diversity. Ephemeroptera species present distinct responses to environmental deterioration such that poses this assemblage between the most useful groups in biomonitoring programs. We hypothesized an alteration in beta diversity at the longitudinal species gradient, which will be influenced mainly by species losses between zones. Changes in temporal beta diversity is also expected, but the main drivers of such alterations will be the species turnover between the sampling period. Ephemeroptera community was monitored in nine sampling points from Itaipu Reservoir, where were installed three sets of substrates composed by a float and 2 wooden substrates. We took biological samples in triplicates monthly, from June-01 to August-02. Our initial hypothesis was partially supported and with significant variations only for spatial approach, between the Reservoir zones. The generated ordering from Non-Metric Dimensional Scale - NMDS - corroborated with spatial analyzes, with the formation of two groups along the gradient zonation of the reservoir. The temporal ordination showed no clear pattern. As expected, the contribution to beta diversity was different for our two approaches, such that the loss of species was more important along the spatial gradient and despite of no significant result, the species replacement was more important among months. The spatial results lead us to infer that differences in limnological characteristics between zones are important for determining differences in Ephemeroptera composition and can reflect the dependency degree of the species in relation to the lentic and sometimes-lotic conditions, mainly in the riverine zone of reservoirs. On the other hand, the absence of a temporal pattern can be result of chaotic variations in the physical and chemical attributes imposed by the reservoir operation, disrupting continuity of the biota and natural succession processes.

Resumo As operações de reservatórios alteram, eliminam ou restringem os ciclos hidrológicos naturais. A comunidade biótica torna-se sujeita a estes eventos não-cíclicos, respondendo com a redução da diversidade de espécies. As espécies de Ephemeroptera apresentam respostas distintas à deterioração ambiental, sendo o grupo muitousado em programas de biomonitoramento. Nós hipotetizamos uma alteração na diversidade beta das espécies no gradiente longitudinal, influenciada principalmente pelo componente aninhamento com a consequente perda de espécies entre as zonas do reservatório. Também são esperadas mudanças na diversidade beta entre os meses de amostragem, mas o principal fator dessas alterações será a substituição das espécies entre os períodos, em função dos rápidos ciclos de vida que o grupo apresenta. A comunidade Ephemeroptera foi monitorada em nove pontos amostrais do reservatório de Itaipu, onde foram instalados três conjuntos de substratos compostos por um flutuador e 2 substratos de madeira. Tomamos amostras biológicas em triplicatas mensalmente, de junho de 2001 a agosto de 2002. Nossa hipótese inicial foi parcialmente suportada, com variações significativas apenas na escala espacial. A ordenação gerada pelo Escalonamento Dimensional Não-Métrico - NMDS - corroborou com as análises espaciais, com a formação de dois grupos ao longo do gradiente de zonação do reservatório. A ordenação temporal não mostrou um padrão claro. Como esperado, a contribuição para a diversidade beta foi diferente para nossas duas abordagens, de modo que a perda de espécies foi mais importante ao longo do gradiente espacial e apesar de não haver resultado significativo, a substituição das espécies foi mais importante entre os meses de amostragem. Os resultados espaciais levam-nos a inferir que as diferenças nas características limnológicas entre as zonas são importantes para determinar as diferenças na composição de Ephemeroptera e podem refletir o grau de dependência das espécies em relação às condições lênticas e às vezes lóticas, principalmente na zona fluvial dos reservatórios. Por outro lado, a ausência de um padrão temporal pode ser resultado de variações caóticas nos atributos físicos e químicos impostos pela operação do reservatório, que interrompem a continuidade da biota e os processos de sucessão naturais.

Reservoir longitudinal gradient promotes ordered losses on diversity and density of Ephemeroptera community.

Reservoir operations alter, eliminate or restrain the natural hydrologic cycles. Biotic community has become subject to these non-cyclic events, responding by reducing the species diversity. Ephemeroptera species present distinct responses to environmental deterioration such that poses this assemblage between the most useful groups in biomonitoring programs. We hypothesized an alteration in beta diversity at the longitudinal species gradient, which will be influenced mainly by species losses between zones. Changes in temporal beta diversity is also expected, but the main drivers of such alterations will be the species turnover between the sampling period. Ephemeroptera community was monitored in nine sampling points from Itaipu Reservoir, where were installed three sets of substrates composed by a float and 2 wooden substrates. We took biological samples in triplicates monthly, from June-01 to August-02. Our initial hypothesis was partially supported and with significant variations only for spatial approach, between the Reservoir zones. The generated ordering from Non-Metric Dimensional Scale - NMDS - corroborated with spatial analyzes, with the formation of two groups along the gradient zonation of the reservoir. The temporal ordination showed no clear pattern. As expected, the contribution to beta diversity was different for our two approaches, such that the loss of species was more important along the spatial gradient and despite of no significant result, the species replacement was more important among months. The spatial results lead us to infer that differences in limnological characteristics between zones are important for determining differences in Ephemeroptera composition and can reflect the dependency degree of the species in relation to the lentic and sometimes-lotic conditions, mainly in the riverine zone of reservoirs. On the other hand, the absence of a temporal pattern can be result of chaotic variations in the physical and chemical attributes imposed by the reservoir operation, disrupting continuity of the biota and natural succession processes.

Effects of food availability and habitat features on the Ephemeroptera species composition at seasonal and spatial scales from neotropical floodplain rivers / Efeitos da disponibilidade de alimento e características do habitat sobre a composição de espécies de Ephemeroptera nas escalas sazonal e espacial de rios de planície neotropicais

Abstract Brazilian floodplains have suffered great changes in their natural characteristics in recent decades, mainly in the flood pulse. The Upper Paraná River floodplain is one of the few places where are found remained areas in which such peculiar characteristics keep reflecting on its high biodiversity. Ephemeroptera nymphs are one of the higher density groups among benthic community, occurring in many water bodies like large rivers and secondary channels. We sought to understand which factors are needed for the species establishment and how much important is the species colonization, especially in environments with anthropogenic changes. The marginal areas, which are more structured with presence of macrophytes, showed the highest density and richness even in the Paraná River that has great human impact. We verified dominance of Americabaetis alphus, Tricorythopsis araponga, Tricorythopsis artigas on the Parana River, correlated with transparency, depth and electric conductivity, while the dominance of Traverella sp. was correlated with water temperature, especially in marginal areas. Consequently, the increasing transparency and electric conductivity due to the Porto Primavera dam in Parana River can be favoring those Ephemeroptera species. We demonstrated the importance of preserving the wetlands of Ivinhema River State Park mainly for Guajirolus sp., which was only registered in this region. Therefore, our study provides support for understanding gaps from previously studies using artificial substrates in three large rivers which are of great importance to the upper Paraná River floodplain.

Resumo Muitas planícies de inundação brasileiras vêm sofrendo enormes alterações em suas características naturais nas últimas décadas, principalmente no pulso de inundação. A planície de inundação do alto rio Paraná é um dos poucos lugares onde ainda há áreas nas quais essas características peculiares se mantêm, o que reflete em sua alta biodiversidade. Dentre a comunidade zoobentônica, Ephemeroptera é um dos grupos com maior densidade, ocorrendo em diversos corpos aquáticos como rios de grande porte e canais secundários. Nós buscamos compreender quais fatores são necessários para o estabelecimento das espécies e o quão importante é a colonização dessas espécies, principalmente em ambientes com modificações antrópicas. As áreas marginais mais estruturadas com presença de macrófitas apresentaram altas densidade e riqueza mesmo no rio Paraná que apresenta grande impacto antrópico. Verificamos que as dominâncias de Americabaetis alphus, Tricorythopsis araponga e Tricorythopsis artigas no rio Paraná foram correlacionadas com maiores valores de transparência, profundidade e condutividade, enquanto que a dominância de Traverella sp. foi correlacionada com temperatura, principalmente em áreas marginais. Consequentemente, o aumento da transparência e da condutividade devido à barragem de Porto Primavera no rio Paraná pode estar favorecendo essas espécies de Ephemeroptera. Nossos resultados demonstram a importância da preservação do Parque nacional das várzeas do rio Ivinhema, principalmente para o gênero Guajirolus sp., o qual apenas foi registrado nessa região. Portanto, nosso estudo vem trazer subsídios para compreensão de lacunas verificadas em estudos anteriores utilizando substratos artificiais em três rios de enorme influência na planície de inundação do alto rio Paraná.

Effects of food availability and habitat features on the Ephemeroptera species composition at seasonal and spatial scales from neotropical floodplain rivers.

Brazilian floodplains have suffered great changes in their natural characteristics in recent decades, mainly in the flood pulse. The Upper Paraná River floodplain is one of the few places where are found remained areas in which such peculiar characteristics keep reflecting on its high biodiversity. Ephemeroptera nymphs are one of the higher density groups among benthic community, occurring in many water bodies like large rivers and secondary channels. We sought to understand which factors are needed for the species establishment and how much important is the species colonization, especially in environments with anthropogenic changes. The marginal areas, which are more structured with presence of macrophytes, showed the highest density and richness even in the Paraná River that has great human impact. We verified dominance of Americabaetis alphus, Tricorythopsis araponga, Tricorythopsis artigas on the Parana River, correlated with transparency, depth and electric conductivity, while the dominance of Traverella sp. was correlated with water temperature, especially in marginal areas. Consequently, the increasing transparency and electric conductivity due to the Porto Primavera dam in Parana River can be favoring those Ephemeroptera species. We demonstrated the importance of preserving the wetlands of Ivinhema River State Park mainly for Guajirolus sp., which was only registered in this region. Therefore, our study provides support for understanding gaps from previously studies using artificial substrates in three large rivers which are of great importance to the upper Paraná River floodplain.

Chironomidae feeding habits in different habitats from a Neotropical floodplain: exploring patterns in aquatic food webs.

Ecological studies on food webs have considerably increased in recent decades, especially in aquatic communities. Because Chironomidae family are highly specious, occurring in almost all aquatic habitats is considered organisms-key to initiate studies on ecological relationships and trophic webs. We tested the hypothesis that the diversity of the morphospecies diet reflects differences on both the food items available among habitats and the preferences of larval feeding. We analyzed the gut content of the seven most abundant Chironomidae morphospecies of the different habitats from the Upper Paraná River. We categorized the food items found into algae, fungal spores, fragments of plants, algae and animal fragments and sponge spicules. We observed the algae predominance in the gut content of morphospecies from lakes. Considering the different regions from each lake, we registered the highest food abundance in the littoral regions in relation to the central regions. From the variety of feeding habits (number of item kinds), we classified Chironomus strenzkei, Tanytarsus sp.1, Procladius sp.1 as generalist morphospecies. We found a nested pattern between food items and Chironomidae morphospecies, where some items were common to all taxa (e.g., Bacillariophyceae algae, especially), while others were found in specific morphospecies (e.g., animals fragments found in Procladius sp.1). The algae represented the most percentage of gut contents of Chironomidae larvae. This was especially true for the individuals from littoral regions, which is probably due to the major densities of algae associated to macrophytes, which are abundant in these regions. Therefore, the feeding behavior of these morphospecies was generalist and not selective, depending only of the available resources.

Limnoperna fortunei Dunker, 1857 larvae in different environments of a Neotropical floodplain: relationships of abiotic variables and phytoplankton with different stages of development.

Limnoperna fortunei Dunker, 1857 is an Asian invasive freshwater bivalve. Although there need to contain their spread, studies about the biology of the larvae are scarce. We correlated the larval stages of L. fortunei with biotic factors such as phytoplankton and main abiotic variables in lotic environments of the Upper Paraná River floodplain. The four samples were taken quarterly during the year 2012. The Principal component analysis (PCA) showed only spatial differences, as did a Canonical Correspondence Analysis (CCA). High densities of larvae were recorded in all samples the Paraná River and Baía River only in December, especially those in their initial stage. In the biovolume of Class of algae, Bacillarophyceae showed the highest value, but Chlorophycea who was strongly correlated with the density of D-stage larvae. The large variety of phytoplankton, especially microplankton Chlorophyceae, high values of PO4, NH4 and temperature were positively correlated with high densities of D-stage larvae. We conclude that high temperature, and food availability, indicated by phytoplankton community, favored the reproduction of L. fortunei and enhance the ability of specie dispersion due to the increase in the emission of propagules. Therefore, studies that address the biology of golden mussel larvae should be performed in order to prevent its spread.

Chironomidae feeding habits in different habitats from a Neotropical floodplain: exploring patterns in aquatic food webs / Hábitos alimentares de Chironomidae em diferentes habitats de uma planície de inundação Neotropical: explorando padrões em redes alimentares aquáticas

Abstract Ecological studies on food webs have considerably increased in recent decades, especially in aquatic communities. Because Chironomidae family are highly specious, occurring in almost all aquatic habitats is considered organisms-key to initiate studies on ecological relationships and trophic webs. We tested the hypothesis that the diversity of the morphospecies diet reflects differences on both the food items available among habitats and the preferences of larval feeding. We analyzed the gut content of the seven most abundant Chironomidae morphospecies of the different habitats from the Upper Paraná River. We categorized the food items found into algae, fungal spores, fragments of plants, algae and animal fragments and sponge spicules. We observed the algae predominance in the gut content of morphospecies from lakes. Considering the different regions from each lake, we registered the highest food abundance in the littoral regions in relation to the central regions. From the variety of feeding habits (number of item kinds), we classified Chironomus strenzkei, Tanytarsus sp.1, Procladius sp.1 as generalist morphospecies. We found a nested pattern between food items and Chironomidae morphospecies, where some items were common to all taxa (e.g., Bacillariophyceae algae, especially), while others were found in specific morphospecies (e.g., animals fragments found in Procladius sp.1). The algae represented the most percentage of gut contents of Chironomidae larvae. This was especially true for the individuals from littoral regions, which is probably due to the major densities of algae associated to macrophytes, which are abundant in these regions. Therefore, the feeding behavior of these morphospecies was generalist and not selective, depending only of the available resources.

Resumo Estudos sobre redes alimentares têm aumentado consideravelmente nas últimas décadas em pesquisas de ecologia, principalmente em comunidades aquáticas. Em função Família Chironomidae ser altamente especiosa, ocorrendo em quase todos os hábitats aquáticos, é considerada como organismos-chave para estudos preliminares sobre as relações ecológicas como redes tróficas. Nós testamos a hipótese de que a diversidade de dietas das morfoespécies reflete diferenças tanto na disponibilidade dos itens alimentares entre os hábitats analisados, quanto preferencias alimentares das larvas. Nós analisamos o conteúdo digestivo das sete morfoespécies mais abundantes de diferentes habitats da planície de inundação do alto rio Paraná. Nós categorizamos os itens alimentares encontrados como algas, esporos de fungos, fragmentos vegetais, fragmentos de algas ou de animais e espículas de esponjas. Nós observamos a predominância de algas no conteúdo digestivo das morfoespécies lacustres. Considerando as diferentes regiões de cada lagoa, nós registramos as maiores abundâncias de alimentos nas regiões litorâneas, quando comparadas com as regiões centrais. A partir da variedade de hábitos alimentares (número de tipos de itens), classificamos Chironomus strenzkei, Tanytarsus sp.1 e Procladius sp.1 como morfoespécies generalistas. Nós encontramos um padrão aninhado entre os itens alimentares e as morfoespécies de Chironomidae, onde alguns itens foram comuns a todos os táxons (por exemplo, algas da Classe Bacillariophyceae), enquanto outros foram encontrados em morfoespécies mais especificas (por exemplo, fragmentos de animais encontrados em Procladius sp.1). As algas representaram a maior porcentagem do conteúdo digestivo das larvas de Chironomidae. Este fato foi especialmente verdadeiro para os indivíduos das regiões litorâneas, o qual é devido, provavelmente, pelas maiores densidades de algas associadas à macrófitas, que são abundantes nessas regiões. Portanto, o comportamento alimentar dessas morfoespécies foi generalista e não seletivo, dependendo apenas dos recursos disponíveis.

Limnoperna fortunei Dunker, 1857 larvae in different environments of a Neotropical floodplain: relationships of abiotic variables and phytoplankton with different stages of development / Larvas de Limnoperna fortunei Dunker, 1857 em diferentes ambientes de uma planície de inundação Neotropical: relação com variáveis abióticas e algas fitoplanctônicas em diferentes estágios de desenvolvimento

Abstract Limnoperna fortunei Dunker, 1857 is an Asian invasive freshwater bivalve. Although there need to contain their spread, studies about the biology of the larvae are scarce. We correlated the larval stages of L. fortunei with biotic factors such as phytoplankton and main abiotic variables in lotic environments of the Upper Paraná River floodplain. The four samples were taken quarterly during the year 2012. The Principal component analysis (PCA) showed only spatial differences, as did a Canonical Correspondence Analysis (CCA). High densities of larvae were recorded in all samples the Paraná River and Baía River only in December, especially those in their initial stage. In the biovolume of Class of algae, Bacillarophyceae showed the highest value, but Chlorophycea who was strongly correlated with the density of D-stage larvae. The large variety of phytoplankton, especially microplankton Chlorophyceae, high values of PO4, NH4 and temperature were positively correlated with high densities of D-stage larvae. We conclude that high temperature, and food availability, indicated by phytoplankton community, favored the reproduction of L. fortunei and enhance the ability of specie dispersion due to the increase in the emission of propagules. Therefore, studies that address the biology of golden mussel larvae should be performed in order to prevent its spread.

Resumo Limnoperna fortunei Dunker, 1857 é um bivalve asiático dulcícola invasor. Embora haja necessidade de conter sua dispersão, estudos que abordam a biologia de suas larvas ainda são escassos. Analisou-se as fases larvais de L. fortunei relacionando-as a fatores bióticos como a comunidade fitoplânctônica e às principais variáveis abióticas, em ambientes lóticos da planície de inundação do alto do rio Paraná. As quatro coletas foram trimestrais durante o ano de 2012. A análise de componentes principais (PCA) demonstrou apenas diferenças espaciais, assim como a Análise de Correspondência Canônica (CCA). Altas densidades de larvas foram registradas em todas as coletas no rio Paraná e no rio Baía apenas no mês de dezembro, principalmente para as larvas em estádio inicial. No biovolume das classes de algas, Bacillarophyceae obteve o maior valor, porém Chlorophycea foi a que fortemente correlacionou-se com a densidade de larvas D. A grande variedade de fitoplâncton, em especial de Chlorophyceae microplanctônica, altos valores de PO4, NH4 e temperatura estiveram positivamente correlacionadas com altas densidades de larvas D. Conclui-se que, altas temperaturas, e disponibilidade de alimento, como a comunidade fitoplanctônica, favorecem a reprodução de L. fortunei e aumentam a capacidade de dispersão da espécie devido ao incremento na emissão de propágulos. Portanto, estudos que abordem a biologia das larvas de mexilhão-dourado devem ser realizados a fim de evitar sua propagação.

Dero (Allodero) lutzi Michaelsen, 1926 (Oligochaeta: Naididae) associated with Scinax fuscovarius (Lutz, 1925) (Anura: Hylidae) from Semi-deciduous Atlantic Rain Forest, southern Brazil.

Amphibians are hosts for a wide variety of ecto- and endoparasites, such as protozoans and parasitic worms. Naididae is a family of Oligochaeta whose species live on a wide range of substrates, including mollusks, aquatic macrophytes, sponges, mosses, liverworts, and filamentous algae. However, some species are known as endoparasitic from vertebrates, such as Dero (Allodero) lutzi, which is parasitic of the urinary tracts of frogs, but also have a free-living stage. Specimens in the parasitic stage lack dorsal setae, branchial fossa, and gills. Here we report the occurrence of D. (A.) lutzi associated with anuran Scinax fuscovarius from Semi-deciduous Atlantic Rain Forest in southern Brazil. The study took place at the Caiuá Ecological Station, Diamante do Norte, Paraná, southern Brazil. Seven specimens of S. fuscovarius were examined for parasites but only one was infected. Parasites occurred in ureters and urinary bladder. Previous records of this D. (A.) lutzi include the Brazilian States of Santa Catarina, São Paulo, Rio de Janeiro, and Minas Gerais, as well as Cuba and North America. This is a new locality record for this species in Brazil. Reports of Dero (Allodero) lutzi are rare, due to difficulty of observation, and such events are restricted only the fortuitous cases. It is important to emphasize the necessity of future studies, which are fundamental to the understanding of biological and ecological aspects of this species.

Chironomidae larvae (Diptera) of Neotropical floodplain: overlap niche in different habitats.

The niche overlap between trophic groups of Chironomidae larvae in different habitats was observed between trophic groups and between different environments in Neotropical floodplain. For the evaluation we used the index of niche overlap (CXY) and analysis of trophic networks, both from the types and amount of food items identified in the larval alimentary canal. In all environments, the larvae fed on mainly organic matter such as plants fragments and algae, but there were many omnivore larvae. Species that have high values of food items occurred in diverse environments as generalists with great overlap niche and those with a low amount of food items with less overlap niche were classified as specialists. The largest number of trophic niche overlap was observed among collector-gatherers in connected floodplain lakes. The lower values of index niche overlap were predators. The similarity in the diet of different taxa in the same niche does not necessarily imply competition between them, but coexistence when the food resource is not scarce in the environment even in partially overlapping niches.

Tributaries as richness source for Oligochaeta assemblage (Annelida) of Neotropical dammed river.

Tributaries may serve as richness source for the river main channel and the zoobenthos community is a good tool to verify this kind of pattern. In this study, we aimed to characterize the benthic invertebrate assemblage in three tributaries associated to the Paraná River main channel, focusing in Oligochaeta community. We hypothesized that (i) in tributaries, Oligochaeta are richer than the main river (Paraná River) and (ii) dammed tributary (Paranapanema River) is poorly diverse than the others. Samples were conducted in Paranapanema, Baía and Ivinhema tributaries using a modified Petersen grab along three transects (samples conducted inside the tributary, in the mouth of each tributary and inside Paraná River). To analyze (i) the difference between the richness and density among the tributaries and the Paraná River and (ii) effect of each tributary transect on the Oligochaeta richness we used a nonparametric Kruskal-Wallis test. Changes in environmental variables and in richness and composition of Oligochaeta were summarized by Canonic Correspondence Analysis. It was registered 21 different benthic invertebrates taxa, being Oligochaeta assemblage with the highest density. Within Oligochaeta, Narapa bonettoi was the most abundant species, followed by Haplotaxis aedochaeta and Paranadrilus descolei. In our results we refused both hypotheses, because we did not found significant differences for richness and density between the tributaries and the main river, and also no difference between the three transects of each tributary were found. However, the tributaries less influenced by damming, especially the Baía recorded high richness. This corroborates their importance to diversity in the floodplain and the species of Oligochaeta reflect the peculiar characteristics of habitats within each tributaries.