Abstract The aquatic insects of the genus Phylloicus play a crucial role in aquatic ecosystems, shredding leaves and contributing to nutrient cycling in streams. Therefore, this genus is often used in laboratory experiments. However, in Cerrado regions such as Araguaia, these studies are impractical due to a lack of knowledge about basic aspects, such as their feeding preferences for local plants. Hence, our objective is to determine the native plant species in the Araguaia region preferred as food by Phylloicus. We conducted an experimental study comparing the consumption of three native Cerrado plant species: Casearia sylvestris, Astronium fraxinifolium, and Ficus guaranítica by Phylloicus. To assess differences in consumption, we performed an Analysis of Variance. The results revealed that Phylloicus larvae exhibited a feeding preference for Casearia sylvestris (F = 9.71; p = 0.004). This finding will contribute to the development of future experimental studies using Phylloicus in the Araguaia region, as understanding the feeding preferences of animals used in experiments is essential for their maintenance in the laboratory. ecosystems streams Therefore However aspects plants Hence fraxinifolium Variance F 9.71 971 9 71 0.004. 0004 0.004 . 0 004 0.004) 9.7 97 7 000 0.00 00 9. 0.0 0.

Resumo Os insetos aquáticos do gênero Phylloicus desempenham um papel essencial nos ecossistemas aquáticos, fragmentando folhas e contribuindo para a ciclagem de nutrientes nos riachos. Por isso, esse gênero é frequentemente utilizado em experimentos de laboratório. No entanto, em regiões de Cerrado como o Araguaia, esses trabalhos são inviáveis por não se conhecer aspectos básicos como a sua preferência alimentar por plantas locais. Por isso, nosso objetivo é responder quais são as espécies vegetais nativas da região do Araguaia preferidas para a alimentação de Phylloicus. Fizemos um trabalho experimental comparando o consumo de três espécies vegetais nativas do Cerrado: Casearia sylvestris, Astronium fraxinifolium e Ficus guaranítica pelos Phylloicus. Para avaliar as diferenças no consumo, realizamos uma Análise de Variância. Os resultados obtidos revelaram que as larvas de Phylloicus demonstraram preferência alimentar por Casearia sylvestris (F = 9.71; p = 0.004). Esse achado ajudará no desenvolvimento de futuros trabalhos experimentais utilizando Phylloicus na região do Araguaia, uma vez que é essencial o conhecimento da preferência alimentar dos animais utilizados nos experimentos para sua manutenção em laboratório. riachos isso laboratório entanto locais Variância F 9.71 971 9 71 0.004. 0004 0.004 . 0 004 0.004) 9.7 97 7 000 0.00 00 9. 0.0 0.

Abstract: Aim We evaluated the effects of native vegetation and agricultural activities on functional characteristics of aquatic insects’ assemblages in Atlantic Forest streams southern Brazil. Methods We collected information on land uses, riparian zone structural characteristics, and limnological variables of ten streams to characterize their environmental quality. In the same streams, we collected aquatic insects (Ephemeroptera, Plecoptera, and Trichoptera) and determined their functional characteristics (e.g. food habits, mobility, shape, and body size). The information for each trait category was quantified using a fuzzy code technique. To test the environmental conditions on aquatic insect functional attributes, we used an RLQ analysis. Results In our study, only five environmental variables showed variation in streams (temperature, pH, DO, native vegetation, and agriculture use). We collected a total of 2591 organisms distributed in 21 EPT genera. We observed that the functional attributes of EPT formed distinct groups in relation to the stream categories defined by the measured environmental variables. We observed that shredders and larger and flattened organisms were associated with natural streams. These attributes were associated with the genera Phylloicus, Anacroneuria, Tupiperla, and Farrodes. On the other hand, we observed greater proportions of scrapers, with spherical bodies and shelter builders in streams impacted by agriculture. The most frequent genera in these streams were Itaura, Wormaldia and Helicopsyche. Conclusions The modification of natural landscapes by agricultural areas caused significant functional changes in the aquatic invertebrate communities. Since aquatic insects participate effectively in ecological processes, alterations in the functional characteristics of these communities can cause changes in the streams’ functioning. Abstract Brazil uses quality Ephemeroptera, Ephemeroptera (Ephemeroptera Plecoptera Trichoptera e.g. eg e g (e.g habits mobility shape size. size . size) technique analysis study temperature, temperature (temperature pH DO use. use use) 259 2 Phylloicus Anacroneuria Tupiperla Farrodes hand scrapers Itaura Helicopsyche processes functioning e.g 25

Resumo: Objetivo Neste estudo avaliamos os efeitos da vegetação nativa e atividades agrícolas sobre as características funcionais de assembleias de insetos aquáticos em riachos de Mata Atlântica, sul do Brasil. Métodos Coletamos informações sobre usos do solo, características estruturais da zona ripária e variáveis limnológicas de dez córregos para caracterizar sua qualidade ambiental. Nos mesmos riachos coletamos insetos aquáticos (Ephemeroptera, Plecoptera e Trichoptera) e determinamos as características funcionais (e.g., hábitos alimentares, mobilidade, forma, tamanho do corpo). A informação para cada categoria de característica foi quantificada usando uma técnica de código fuzzy. Para testar as condições ambientais nos atributos funcionais de insetos aquáticos, utilizamos uma análise RLQ. Resultados Em nosso estudo, apenas cinco variáveis ambientais apresentaram variação entre os riachos (temperatura, pH, DO, vegetação nativa, uso agrícola). Coletamos um total de 2591 organismos distribuídos em 21 gêneros das ordens EPT. Observamos que os atributos funcionais do EPT formaram grupos distintos em relação às categorias de riachos definidas pelas medidas das variáveis ambientais. Observamos que fragmentadores e organismos maiores e achatados estiveram associados a riachos naturais. Esses atributos foram associados aos gêneros Phylloicus, Anacroneuria, Tupiperla e Farrodes. Por outro lado, em riachos impactados pela agricultura, observamos maiores proporções de raspadores, com corpos esféricos e construtores de abrigos. Nesses riachos, os gêneros mais frequentes foram Itaura, Wormaldia e Helicopsyche. Conclusões A modificação das paisagens naturais por áreas agrícolas causou mudanças funcionais significativas nas comunidades de invertebrados aquáticos. Como os insetos aquáticos participam efetivamente dos processos ecológicos, alterações nas características funcionais dessas comunidades podem causar mudanças no funcionamento dos riachos. Resumo Atlântica Brasil solo ambiental Ephemeroptera, Ephemeroptera (Ephemeroptera Trichoptera e.g., eg g (e.g. alimentares mobilidade forma corpo. corpo . corpo) fuzzy RLQ temperatura, temperatura (temperatura pH DO agrícola. agrícola agrícola) 259 2 Phylloicus Anacroneuria Farrodes lado agricultura raspadores abrigos Itaura Helicopsyche ecológicos e.g. (e.g 25 e.g

RESUMO Nós criamos massas de ovos dos tricópteros Phylloicus amazonas (n = 2) e Phylloicus elektoros (n = 7) coletadas no campo até a fase adulta. A mediana do tempo de desenvolvimento do primeiro instar larval ao adulto foi de 229 dias para P. amazonas e 275 dias para P. elektoros, incluindo o tempo de desenvolvimento pupal (mediana de 13.5 e 16 dias, respectivamente). Fragmentadores têm sido frequentemente usados em estudos relacionados ao processamento de matéria orgânica em ecossistemas aquáticos. As informações biológicas fornecidas neste estudo podem ser úteis para o planejamento de estudos experimentais em campo ou em condições de laboratório.

ABSTRACT We reared field-collected egg masses of the caddisflies Phylloicus amazonas (n = 2) and Phylloicus elektoros (n = 7) to adulthood. First instar larvae of both species hatched up to five days after field collection. Median development time from first larval instar to adult was 229 days for P. amazonas and 275 for P. elektoros, including the pupal development time (median of 13.5 and 16 days, respectively). Shredder organisms have been used frequently in studies related to organic matter processing in freshwater ecosystems. The biological information reported in here may be useful for the design of experimental studies in the field or under laboratory conditions.

Resumo: Objetivo Avaliar a situação geral dos estudos sobre a decomposição da matéria orgânica vegetal em ambientes aquáticos continentais da região subtropical da América do Sul. Métodos Utilizamos bancos de dados e compilamos publicações (1990-2018) utilizando combinações de palavras referentes à decomposição, detritos, ambientes aquáticos e países da América do Sul com clima subtropical. As características das publicações foram analisadas ao longo do tempo. Resultados Um total de 1.042 publicações foram encontradas, com 106 tratando especificamente do objetivo deste estudo. Em clima subtropical na América do Sul este tópico de pesquisa é incipiente. A maioria dos estudos foi realizada em riachos (39,6%) e avaliou o processo de decomposição e a associação de invertebrados (42,5%). Há uma grande deficiência em estudos realizados em ambientes lênticos (e.g. lagos, 10%). Conclusão Os estudos sobre a decomposição de detritos orgânicos em ecossistemas aquáticos na região subtropical da América do Sul são ainda emergentes. Enquanto o conhecimento sobre o papel dos fragmentadores é razoável, poucos estudos têm avaliado a importância de microrganismos no processo de decomposição. Finalmente, nosso estudo demonstrou que as colaborações científicas impulsionaram as publicações sobre o tema.

Abstract: Aim To evaluate the general situation of studies on the decomposition of organic matter in aquatic environments in the subtropical region of South America. Methods We compiled publications (1990-2018) with a combination of words referring to decomposition, debris, aquatic environments and South American countries with subtropical climate. The characteristics of the publications were analysed over time. Results A total of 1,042 publications were found, with 106 specifically addressing the purpose of this study. For subtropical climates in South America, this research topic is incipient. Most studies were carried out in streams (39.6%) and studied the decomposition process and associated invertebrates (42.5%). There is a great deficiency of studies in lentic environments (e.g. lakes, 10%). Conclusion Studies on the decomposition of organic detritus in aquatic ecosystems in the subtropical region of South America are still emerging. While the knowledge about the role of shredders is more or less understood, the importance of microorganisms on the decomposition process still needs to be better evaluated. Finally, our study demonstrated that scientific collaboration boosted publication on the topic.

Abstract Investigations on the fungal community associated with the digestive tract (DT) of insects have provided insights into the diversity of associated microorganisms and their potential roles in the interaction with their hosts. However, most studies have focused on terrestrial insects, with few studies focusing on aquatic insects in Neotropical regions. We studied fungal taxa associated with the DT of larval stages of the aquatic shredders Phylloicus amazonas, P. elektoros and P. fenestratus in the Brazilian Amazon Forest. Filamentous fungi were isolated, purified and screened for cellulolytic activity. A total of 33 fungal taxa was identified through the combination of classical and molecular taxonomy. The genus Penicillium was the most frequent in DT of Phylloicus spp. (18.75%). The occurrence of fungal taxa among hosts was quite variable, with more than half of the associated fungi being exclusive of each host species. A significant portion of the fungal community associated with each host presented cellulolytic activity (± 50%). It was concluded that the fungal community associated with Phylloicus spp. larvae consist mainly of fungal taxa from food items, which come from riparian vegetation (whose plant species are variable) or are indigenous of the aquatic ecosystems, which is the habitat of these larvae.

RESUMO Fungos filamentosos tem sido alvo de estudos de bioprospecção devido à sua grande eficiencia em produzir enzimas extracelulares, as quais tem grande potencial para uso em bioindústrias. Neste estudo, fungos filamentosos foram isolados do intestino de larvas de insetos aquáticos da Amazônia, para avaliar sua atividade celulolítica. Foram coletadas 69 larvas de insetos aquáticos fragmentadores de três gêneros: Phylloicus (Trichoptera: Calamoceratidae), Triplectides (Trichoptera: Leptoceridae) e Stenochironomus (Diptera: Chironomidae) em dez igarapés de uma área protegida na Amazônia central brasileira. O crescimento dos fungos isolados foi feito em meio de cultura Ágar Batata Dextrose (BDA). Os isolados fúngicos foram transferidos para o meio sintético com Carboximetil celulose e utilizou-se vermelho Congo para determinar o índice enzimático. O halo de hidrólise, indicando a produção de celulases, foi observado em 175 isolados fúngicos (70% do total), dos quais 25 tiveram um índice enzimático ≥ 2,0 e pertencem a sete gêneros fúngicos. Os táxons fúngicos Cladosporium, Gliocephalotrichum, Penicillium, Pestalotiopsis, Talaromyces, Trichoderma e Umbelopsis foram isolados dos intestinos das larvas de Phylloicus, Triplectides e Stenochironomus e são tradicionalmente utilizados em aplicações biotecnológicas. Os resultados indicam um potencial celulolítco destes fungos associados ao intestino de insetos aquáticos amazônicos.

ABSTRACT Filamentous fungi have been targeted by bioprospecting studies because they are effective producers of extracellular enzymes that can potentially be used by the bioindustry. In this study, we isolated filamentous fungi from the guts of Amazonian aquatic insect larvae to evaluate their cellulolytic activity. We collected 69 larvae of shredder insects of three genera: Phylloicus (Trichoptera: Calamoceratidae), Triplectides (Trichoptera: Leptoceridae) and Stenochironomus (Diptera: Chironomidae) in ten streams from a protected area in the central Brazilian Amazon. Production of mycelia was elaborated in PDA (Potato Dextrose Agar) medium. The isolates were transferred to a synthetic medium with carboxymethyl cellulose, and Congo red was used to determine the enzymatic index. The hydrolysis halo, indicating the production of cellulases, was observed in 175 fungal isolates (70% of the total), of which 25 had an enzymatic index ≥ 2.0 and belonged to seven fungal genera. The fungal taxa Cladosporium, Gliocephalotrichum, Penicillium, Pestalotiopsis, Talaromyces, Trichoderma and Umbelopsis were isolated from guts of Phylloicus, Triplectides and Stenochironomus, which are traditionally used in biotechnological applications. Our results indicate the cellulolytic potential of fungi associated with the guts of aquatic Amazonian insects.

Resumen Introducción: En los sistemas fluviales colombianos ha sido poco estudiada la fragmentación del material alóctono por parte de los insectos acuáticos. Uno de los organismos trituradores más abundantes en las corrientes tropicales de América es el tricóptero del género Phylloicus (Calamoceratidae). Objetivo: El presente estudio tuvo como objetivo evaluar el efecto de los atributos químicos (nitrógeno, fósforo y lignina) y físicos (dureza) de las hojas de tres especies ribereñas (Ficus tonduzii, Zygia longifolia y Clusia multiflora) dominantes en la parte media del río Gaira, sobre la preferencia de hojas y el crecimiento de larvas de Phylloicus sp. Métodos: Se realizaron experimentos de multiselección de alimento en campo, en los que se ubicaron cámaras experimentales dentro del río por 21 días. Para la valoración de la calidad foliar se recolectaron hojas del río, las cuales se secaron y se pulverizaron para realizar los análisis químicos. Resultados: No se presentaron diferencias significativas en los porcentajes de lignina y fósforo entre las hojas de las tres especies de árboles, pero si en su dureza y en el contenido de nitrógeno. Las hojas de C. multiflora fueron más suaves (180.1± 53.9 g) que las de F. tonduzii (285.3 ± 88.4 g) y Z. longifolia (232.3 ± 60.8 g), pero sin diferencias entre las dos últimas especies. Las hojas de Z. longifolia también tuvieron más nitrógeno (1.9 ± 0.0%) que las de hojas de las otras especies (1 ± 0.0 %). En cuanto a la preferencia de hojas, las larvas de Phylloicus sp. utilizaron en mayor proporción las hojas de C. multiflora. Aunque las hojas de C. multiflora no tuvieron diferencias significativas en el contenido de lignina y fósforo, presentaron valores menores de dureza, lo que podría hacerlas más palatables para Phylloicus sp. Por otra parte, el crecimiento específico diario de las larvas fue mayor cuando usaron hojas de F. tonduzii. Conclusiones: Nuestros resultados parecen indicar que las hojas más preferidas no son necesariamente las de mejor calidad nutricional y sugieren que la dureza de las hojas es la variable más importante en la selección de la hojarasca por parte de las larvas de Phylloicus sp.

Abstract Introduction: Fragmentation of allochthonous material by aquatic insects is a functional process that has been little studied in Colombian rivers. One of the most abundant shredder genus in American tropical stream ecosystems is the caddisfly genus Phylloicus (Calamoceratidae). Objective: In this study we evaluate the effect of chemical (nitrogen, phosphorus and lignin) and physical (toughness) attributes of leaves of three dominant tree species (Ficus tonduzii, Zygia longifolia and Clusia multiflora) in the riverine area of the middle section of the Gaira river, on leaf preferences and larval growth of a species of Phylloicus. Methods: Multiple food selection field experiments were performed, in which experimental chambers were located in the river for twenty-one days. For the assessment of leaf quality, leaves were collected in the river, dried and pulverized to perform chemical analyzes. Results: There were no significant differences in lignin and phosphorous percentages between the leaves of the three species, but nitrogen content and leaf toughness differed among them. The leaves of C. multiflora were softer (180.1± 53.9 g) than those of F. tonduzii (285.3 ± 88.4 g) and Z. longifolia (232.3 ± 60.8 g), but without differences between the last two species. The Z. longifolia leaves also had more nitrogen (1.9 ± 0.0 %) than those of the other species (1 ± 0.0 %). Regarding leaf selection, Phylloicus sp. larvae used in greater proportion the leaves of C. multiflora. Although lignin and phosphorous differences were not significant, the leaves of C. multiflora had lower toughness, which could make them more palatable for Phylloicus sp. On the other hand, the daily specific growth of the larvae was higher when they used leaves of F. tonduzii. Conclusions: Our findings seem to indicate that the most preferred leaves were not necessarily those of better nutritional quality and suggest that leaf toughness was the most important variable in the selection of leaf litter by the larvae of Phylloicus sp.

Abstract Freshwater crabs are macroconsumers that are commonly found in Neotropical headwater streams that may play a key role in energy flow and nutrient cycling in detrital food webs. Although studies have examined the feeding habits of trichodactylid crabs, little is known of this behavior in pseudothelphusid species, and specifically whether they actually consume leaf material. We conducted three nine-day laboratory trials with pseudothelphusid crabs (Ptychophallus tumimanus (Rathbun, 1898)) and leaves (Koanophyllon pittieri) to investigate whether crabs shred leaves. We hypothesized that leaf mass loss would be faster with crabs present relative to control tanks with only leaves. Leaf mass loss was significantly higher (p < 0.001) in tanks with crabs (0.49 ± 0.07 g, mean ± 1 SD) compared to control tanks (0.31 ± 0.05 g). We observed crabs manipulating, shredding, and consuming leaves, with leaf fragments and egesta present in tanks with crabs but not in control tanks. Their consumption and egestion activity may affect nutrient availability and transformation by stimulating microbial activity during leaf breakdown and converting coarse particulate organic matter (CPOM) to fine particulate organic matter (FPOM). Therefore, freshwater crabs need to be considered when studying energy flow and nutrient cycling in detrital food webs of Neotropical headwater streams.

Resumo Macroinvertebrados fragmentadores consomem folhas preferencialmente condicionadas por fungos e bactérias que lhes oferece uma maior palatabilidade. Nas espécies do cerrado esse condicionamento está também associado às altas concentrações de elementos químicos limitantes à palatabilidade como alto teor de lignina e celulose, que tornam as folhas menos atrativas para os fragmentadores. O trabalho teve como objetivo avaliar a preferência alimentar de macroinvertebrados fragmentador ( Phylloicus sp.) por material vegetal de dois diferentes biomas (Cerrado e Mata Atlântica), após condicionamento em riacho de Mata Atlântica, observando suas características físicas e químicas. Foram coletadas folhas senescentes do aporte vegetal (AV) de espécies nativas da vegetação ripária nesses dois biomas, com periodicidade mensal em um trecho de 500 m de um córrego em cada bioma. O experimento foi delineado com dois tratamentos. O primeiro (T1) compreendeu discos de folhas do Cerrado (Chrysophyllum oliviforme) mais discos de folhas da Mata Atlântica (Miconia chartacea) que foram condicionadas em córrego de Mata Atlântica. O segundo tratamento (T2) envolveu discos de folhas da Mata Atlântica condicionadas em córrego da Mata Atlântica mais discos de folhas da Mata Atlântica condicionadas em córrego do Cerrado. Os dois testes apontaram diferenças significativas entre os dois tratamentos (T1 e T2). Para T1 houve consumo de discos de folha de M. chartacea por Phylloicus sp, mas não houve consumo dos discos de C. oliviforme , de Cerrado. Para T2, houve o consumo, porém a preferência pelas folhas de M. chartaceae condicionadas no córrego da Mata Atlantica foi consideravelmete maior. Os resultados apontam que Phylloicus sp, apresentou preferência alimentar pelo detrito de bioma de Mata Atlântica e rejeição pelo detrito do bioma Cerrado.

Abstract Macroinvertebrate shredders consume preferably leaves conditioned by fungi and bacteria which offer greater palatability to them. Plant species in Cerrado present high concentration of chemical elements such as lignin and cellulose, phenols and tanins thus making them less attractive for shredders consumption and limiting the palatability. This study aimed to evaluate the feeding preference of a macroinvertebrate shredder of the genus Phylloicus for plant material from two different biomes (Cerrado and Mata Atlântica), after conditioning in a stream of Mata Atlântica and observing their physical and chemical characteristics. Senescent leaves were collected, monthly from the litterfall of riparian vegetation in a 500 m stretch of a stream in each biome from August 2014 to January 201. The most abundant species in each stream was selected for the experiment. The experimental design consisted in with two treatments. The first (T1) comprised leaf discs from Chrysophyllum oliviforme (Cerrado species) together with leaf discs of Miconia chartacea (Atlantic Forest species) which were conditioned in the Atlantic Forest stream. The second treatment (T2) involved leaf discs of Miconia chartacea conditioned in Mata Altlântica and Cerrado streams. Both tests had showed significant differences between the two treatments (T1 and T2). For T1, there was consumption of M. chartacea leaf discs by Phylloicus sp., but there was no consumption of C. oliviforme discs. For T2, there was preference for M. chartaceae leaves conditioned in a stream of Mata Atlântica than in Cerrado stream. The results showed that Phylloicus sp., had presented preference for food detritus of the Mata Antlântica biome and rejection to the one from Cerrado biome.

Resumen Objetivos: Analizar la sucesión de insectos asociados a la descomposición de cuerpos en dos ecosistemas acuáticos en la Amazonia Andina. Metodología: Tres cadáveres fueron ubicados en una quebrada y tres en un lago, los muestreos se realizaron cada cinco horas desde el momento de la muerte hasta su esqueletización. Resultados: La descomposición total duró 14 días en la quebrada y 16 en el lago. Seis estadios de descomposición fueron observados: fresco sumergido; flotación temprana; flotación tardía; deterioro de la flotación; restos flotantes y restos sumergidos. En total se colectaron 1172 individuos en la quebrada y 1585 en el lago. Los gremios tróficos identificados fueron: trituradores; colectores; depredadores; necrófagos; sarcosaprófagos e incidentales. En la quebrada, los trituradores tricópteros fueron los más representativos en la etapa fresco sumergido. En el lago los trituradores, moscas de mayo y los depredadores hemípteros y odonatos fueron los más representativos en la etapa de flotación temprana. Los dípteros necrófagos y sarcosaprófagos fueron los más representativos en las etapas donde el cuerpo estuvo expuesto a la superficie en ambos ecosistemas. Conclusiones: El ISMP para ambos cadáveres fue similar en los dos hábitats, cada uno evidenció seis etapas de descomposición, lo que confirma que la descomposición en los ecosistemas acuáticos es diferente a la observada para los entornos terrestres.

Abstract Objectives: Analyze the succession of insects associated with the decomposition of bodies in two aquatic ecosystems in the Andean Amazon. Scope: Provide information on the insects associated with bodies that decompose within aquatic habitats and the duration times of each decomposition phase as a tool for the determination of the post-mortem submerged interval (PMSI). Methodology: Three carcasses were placed in a stream and three in a lake, the sampling carried out every five hours, from the moment of death to its skeletonization. Main results: Total decomposition lasted for 14 days in the stream and 16 days in the lake. Six decomposition stages were observed: submerged fresh, early floating, floating decay, bloated deterioration, floating remains and sunken remains. A total of 1172 organisms were collected on the carcasses in the stream and 1585 in the lake. Trophic relationships identified were: Shredders, collectors, predators, necrophages, sarcosaprophages and incidentals. In the stream, the caddisfly shredders were the most representative organisms in the submerged fresh stage. In the lake, the mayfly shredders, and hemipterous and odonate predators, were the most representative organisms in the early floating stage. The necrophagous and sarcosaprophagous diptera were the most representative organisms in the stages exposed to the surface in both ecosystems. Conclusions: The PMSI for both carcasses was similar in the two habitats, each showing six stages of decomposition, confirming that decomposition in lotic and lentic ecosystems is different from that observed for terrestrial environments.

RESUMO Nós investigamos em experimento de campo e de laboratório os efeitos das características foliares e condicionamento microbiano de duas espécies de árvores na abundância, biomassa e preferência alimentar de um fragmentador típico. No experimento de campo, comparamos a associação do Phylloicus (Calamoceratidae, Trichoptera) entre duas espécies vegetais, comumente encontradas em zonas ripárias, que são representativas de alta e baixa qualidade nutricional, respectivamente: Nectandra megapotamica e Chusquea tenella. Em experimento de laboratório, investigamos a preferência alimentar de Phylloicus, usando folhas condicionadas e não condicionadas pelos fungos aquáticos. As mesmas espécies vegetais usadas no experimento de campo foram usadas em laboratório. Inicialmente, as folhas de C. tenella mostraram-se mais nutritivas e macias, enquanto que as de N. megapotamica foram mais duras e lignificadas. Os fragmentadores preferiram o detrito com menor dureza e condicionados pelos fungos (campo: C. tenella; laboratório: N. megapotamica, ambas condicionadas por 14 dias), ou seja, estes parecem ser os fatores determinantes para a escolha dos Phylloicus. Após 14 dias, as folhas de N. megapotamica apresentaram decréscimo na dureza, que está associado ao condicionamento microbiano, e que explicou o padrão de consumo por Phylloicus. Nós encontramos evidências em campo e em laboratório que os Phylloicus são seletivos em relação ao alimento e que as características foliares, especialmente relacionadas à estrutura da folha (ex. dureza foliar e lignina), determinam a associação e a preferência destes fragmentadores.

ABSTRACT Through field and laboratory experiments we investigated the effects of leaf traits of two tree species and microbial conditioning on the abundance, biomass, and feeding preference of a typical macroinvertebrate shredder. In the field, we compared the association of Phylloicus (Calamoceratidae, Trichoptera) with two tree species commonly found in riparian zones, which are representative of high and low nutritional quality, respectively: Nectandra megapotamica and Chusquea tenella. In the laboratory, we investigated the feeding preference of Phylloicus using unconditioned leaves and leaves conditioned by aquatic fungi. The same tree species used in the field experiment were used in the laboratory. Initially, C. tenella leaves were proved to be more nutritious and softer, while N. megapotamica leaves were harder and more lignified. The shredders preferred conditioned leaf detritus of reduced toughness (field: C. tenella; laboratory: N. megapotamica, both conditioned for 14 days). These leaf traits seem to be crucial for the choice process of Phylloicus. After 14 days, N. megapotamica leaves showed a decreased toughness associated with the microbial conditioning, which explained its consumption rate by Phylloicus. In both field and laboratory experiments, we found evidence that Phylloicus is a selective feeding shredder, and that the leaf traits, especially leaf structure (e.g., leaf toughness and lignin content), determine its association and preferences.

The aim of this study was to determine if the rate of degradation and the habitat type associated with two submersed macrophytes affect the structure of the invertebrate communities in a shallow subtropical lake. We evaluated debris decomposition rate with a litter bags approach, assigned to four treatments: Potamogeton pectinatus decomposing inside their own stand (PP) and inside a Chara zeylanica stand (PC), and C. zeylanica decomposing inside their own stand (CC) and inside a P. pectinatus stand (CP). During the degradation experiment (20 days), we evaluated the fauna associated both with debris and at live macrophytes. The debris were washed, dried and the coefficient of degradation was determined. We estimated the richness and abundance of taxa of associated invertebrates, and they were classified into functional feeding groups. We evaluated preference of invertebrate communities comparing fauna at live macrophytes and their debris. We observed differences in mass loss between the treatments. Chara zeylanica showed a mass loss four times faster than P. pectinatus. The highest invertebrate abundance occurred in treatment PP and highest invertebrate richness in treatments CC and PP. Collectors and predators showed the highest abundances. The PP treatment had the greatest number of collectors and PC an equitable distribution of all functional groups. Five taxa showed preference for P. pectinatus debris instead of C. zeylanica or P. pectinatus alive, and debris of charophyte was preferred by six taxa instead of alive plant, and by two taxa when the debris were incubated in the stand of P. pectinatus. Our study demonstrated the interference of the incubation site on the decomposition coefficient and on the structuring of the invertebrate communities, and that the abundance of invertebrate at detritus is mainly due to abundance at live plants. Also, the absence of shredders suggests the use of debris mainly as shelter instead of food resource.

El objetivo de este estudio fue determinar si el tiempo de degradación y el hábitat provisto por dos macrófitas sumergidas afectan la estructura de la comunidad de invertebrados en un lago somero subtropical. Se evaluó la descomposición de detrito en 60 bolsas. Los detritos fueron asignados al azar a cuatro tratamientos: Potamogeton pectinatus en descomposición dentro de su propio stand (PP) y en el interior de un stand de Chara zeylanica (PC), y C. zeylanica dentro de su propio stand (CC) y en el interior de un stand de P. pectinatus (CP). Durante el experimento de degradación (20 días) se evaluó la fauna asociada tanto a los detritos como a las macrófitas vivas. Las bolsas de detrito se lavaron, se secaron y luego se determinó el coeficiente de degradación. Estimamos la riqueza y la abundancia de los taxa de invertebrados asociados y evaluamos la preferencia de las comunidades de invertebrados comparando la fauna de las macrófitas vivas y de sus detritos. Los invertebrados asociados se clasificaron en grupos funcionales de alimentación. Chara zeylanica mostró una pérdida de peso cuatro veces más rápida que P. pectinatus. La mayor abundancia de invertebrados se produjo en el tratamiento PP y la mayor riqueza de invertebrados en los tratamientos de CC y PP. Los colectores y depredadores presentaron las mayores abundancias. El tratamiento PP tuvo el mayor número de colectores y PC una distribución más equitativa de los grupos funcionales. Cinco taxones mostraron preferencia por detritos de P. pectinatus en lugar de C. zeylanica o P. pectinatus vivos, y los detritos de la carófita fueron preferidos por seis taxones dentro de las plantas vivas, y por dos taxones cuando se incubaron en el interior del stand de P. pectinatus. Nuestro estudio demostró el efecto del sitio de incubación en el coeficiente de descomposición y en la estructura de las comunidades de invertebrados, y reveló que la abundancia de invertebrados en detritus se debe principalmente a su abundancia en las plantas vivas. Además, la ausencia de trituradores sugiere el uso de detritos principalmente como refugio en lugar de como recurso alimentario.

Resumen Los macroinvertebrados fragmentadores han sido ampliamente reconocidos como un importante grupo funcional alimenticio que contribuye a la descomposición de la hojarasca en quebradas del hemisferio norte, pero poco se sabe sobre su papel en quebradas de montaña neotropicales. En el presente estudio investigamos el efecto de la exclusión de los macroinvertebrados sobre las tasas de fragmentación de la hojarasca en dos quebradas de montaña en Colombia. Nuestra hipótesis es que la fragmentación de la hojarasca es mediada principalmente por macroinvertebrados fragmentadores en quebradas de montaña en Colombia. Este estudio fue llevado a cabo entre enero y abril 2009, incluyendo la época seca y el comienzo de la época de lluvias. Se midieron las pérdidas de masa de hojarasca usando bolsas de malla gruesa (15 mm; accesible a macroinvertebrados) y fina (200 µm; macroinvertebrados excluidos); estas fueron atadas con estacas de madera al fondo de cada quebrada y recuperadas transcurridos 7, 14, 21, 28 y 56 días. Se seleccionó hojarasca de dos especies dominantes en los sitios de estudio y se usaron tres tratamientos para las bolsas de hojarasca: Critoniopsis ursicola, Palicourea cuatrecasasii y una mezcla de estas dos especies. Se compararon las tasas de fragmentación de la hojarasca entre las dos quebradas, los dos tipos de malla y las especies de hojarasca. Los macroinvertebrados que colonizaron las bolsas de hojarasca fueron medidos en términos de densidad, biomasa y categorizados en grupos funcionales alimenticios. En general, nuestros resultados no mostraron diferencias significativas en las tasas de fragmentación de la hojarasca entre las quebradas y entre los tipos de malla. En contraste, las tasas de fragmentación presentaron diferencias significativas entre las especies de hojarasca; específicamente, C. ursicola tuvo tasas más altas de fragmentación que las especies mezcladas y que P. cuatrecasasii. Con respecto a los macroinvertebrados, el presente estudio encontró que su exclusión no tuvo un efecto sobre las tasas de fragmentación y los ensamblajes estuvieron compuestos por una alta densidad de recolectores (principalmente de la familia Chironomidae) y pocos fragmentadores con gran tamaño corporal. La colonización similar de macroinvertebrados entre las especies de hojarasca, las relativamente lentas tasas de fragmentación, sugieren que las hojas fueron un sustrato refractario. Futuros estudios deberían incluir especies de hojarasca con diferentes calidades nutricionales y una mayor escala espacio-temporal para estudiar la hipótesis de la presencia de fragmentadores en quebradas de montaña en Colombia.

Abstract Macroinvertebrate shredders have been widely recognized as an important functional feeding group that contributes to leaf decomposition in temperate streams, but little is known about their role in upland Neotropical streams. In this study, we investigated the effect of macroinvertebrate exclusion on leaf breakdown rates in two upland streams in Colombia. The study was carried out between January and April of 2009, including the dry season and the beginning of the wet season. We measured leaf mass loss using coarse- (15 mm; accessible to macroinvertebrates) and fine- (200 µm; macroinvertebrates excluded) mesh litter bags that were anchored with wood stakes to the bottom of each stream and recovered after 7, 14, 21, 28, and 56 days. We selected leaves from two dominant plant species and used three leaf treatments: two single species (Palicourea cuatrecasasii and Critoniopsis ursicola) and mixed-species (P. cuatrecasasii and C. ursicola). We compared leaf breakdown rates between the Peña Bonita and Marianela stream, mesh-sizes, and leaf treatments. Macroinvertebrates that colonized the litter bags were measured for density and biomass and categorized in functional feeding groups. In general, there were no significant differences in breakdown rates between the streams and the mesh-sizes. In contrast, leaf breakdown rates had significant differences between all leaf species, where C. ursicola had higher breakdown rates than mixed-species, and P. cuatrecasasii. Macroinvertebrate exclusion did not affect leaf breakdown rates and their assemblages were composed by high densities of collectors (mainly Chironomidae) and few shredders with a large body size. The similar macroinvertebrate colonization between leaf species, the relatively slow breakdown rates, and the shredder scarcity, suggest that leaves were a refractory substrate. Further studies should include leaf species with different nutritional qualities and larger spatial-temporal scales to test the hypothesis of shredder presence and its role on leaf decomposition in upland Colombian streams. Rev. Biol. Trop. 66(1): 457-467. Epub 2018 March 01.

RESUMO A atratividade de um determinado detrito foliar pelos invertebrados depende da composição química, estrutura física, estágio de colonização microbiana e nível de degradação do detrito. Além da qualidade do detrito foliar, a temperatura da água tem sido outro fator que pode levar a variações na velocidade da ciclagem de matéria orgânica em ambientes aquáticos. Os objetivos deste estudo foram: i) ampliar os conhecimentos acerca do papel dos invertebrados fragmentadores sobre a atividade de fragmentação de folhas senescentes, senescentes condicionadas e verdes de Richeria grandis Vahl. (Phyllanthaceae) sob elevações graduais de temperatura e verificar as possíveis consequências sobre a perda de massa foliar; ii) verificar a preferência de recurso foliar para a construção de casulo pelo invertebrado fragmentador Phylloicus sp. (Trichoptera). O experimento I foi constituído de três tratamentos que se diferiram apenas na temperatura da água: Tratamento 1: 20°C, Tratamento 2: 24°C, Tratamento 3: 28°C. O experimento II se diferiu do I pelo acréscimo de dez discos de folhas verdes de R. grandis e dois invertebrados (Phylloicus sp.) em cada aquário. A perda de massa foliar na presença ou ausência de atividade de fragmentação não foi influenciada significativamente por diferenças na qualidade intraespecífica do detrito e ou por diferenças na temperatura. A qualidade intraespecífica de recursos misturados com folhas verdes pode levar ao aumento do contato de conteúdos fenólicos sobre folhas senescentes, levando a uma similaridade na exploração alimentar pelos invertebrados. No entanto, a construção de casulos pelos Phylloicus sp. parece ser dependente da resistência apresentada pelo recurso e não do conteúdo fenólico, levando assim os fragmentadores a utilizarem significativamente uma maior quantidade de detritos senescentes, independente da temperatura.

ABSTRACT The attractiveness of a given leaf detritus to invertebrates depends on the chemical composition, physical structure, stage of microbial colonization and decomposition level. In addition to the leaf detritus quality, water temperature also affects organic matter cycling in aquatic environments. In this context, the objectives of this study were: i) to increase knowledge about the role of invertebrate shredders on fragmenting senescent, conditioned senescent and green leaves of Richeria grandis Vahl. (Phyllanthaceae) under gradual temperature rise and check leaf mass loss; ii) to verify the preference of Phylloicus sp. (Trichoptera) for leaves for larval case production. The experiment I consisted of three treatments with different water temperatures: 20°C, 24°C and 28°C. In the experiment II, we added 10 discs of green leaves of R. grandis and two shredder invertebrates (Phylloicus sp.) to each aquarium. Leaf mass loss in the presence or absence of fragmentation was not significantly influenced by differences in the intraspecific quality of the detritus or temperature. Intraspecific quality of resources mixed with green leaves can lead to increased contact of phenolic content on senescent leaves, leading to a similarity in food exploitation by invertebrates. However, the production of larval cases by Phylloicus sp. seems to be dependent on the resistance presented by the resource rather than the phenolic content, thus leading the shredders to use significantly more senescent detritus, regardless temperature.

In small streams, aquatic community obtains energy mainly from leaves of the riparian vegetation. Processing these leaves involves physical, chemical and biological factors that may difer among leaf species. We assessed the efects of litter quality on leaf decomposition and colonization by aquatic invertebrates in a subtropical stream. Leaves from two native, highly represented tree species common in subtropical riparian areas, Ficus luschnathiana and Gasearía sylvestris, were incubated in coarse mesh bags in a stream. Bags were sampled weekly during an incubation of 28 days. The decomposition rates were higher in leaves of F. luschnathiana than in those of C. sylvestris. In the frst seven days of the experiment, leaves of F. luschnathiana lost ~60% of the initial mass, while Cascarla sylvestris lost around 30%. The leaves of C. sylvestris were initially more palatable than the leaves of F. luschnathiana, perhaps due to their lower lignin and cellulose content, and lignin to N ratio (Lignin:N). These structural compounds enhance leaf toughness thereby protecting them from herbivores and detritivores. However, leaves of F. luschnathiana were comparatively softer, which apparently facilitated fungal colonization and conditioning. Thus, leaves of F. luschnathiana attained high shredders density, which was refected in a higher leaf mass loss. Our results suggest that litter decomposition was sensitive to fungal biomass and leaf toughness. We observed that microbial conditioning, rather than the initial chemical quality of the leaf litter, decreased leaf toughness thereby stimulating further colonization by shredders and enhancing the decomposition process. The fungal community has a key-role in the degradation process within aquatic environments, breaking down even low-quality litter (i.e., F. luschnathiana) and promoting changes in the composition and structure of the invertebrate community.

En arroyos de bajo orden, la comunidad acuática obtiene energía principalmente del aporte de la hojarasca que proviene de la vegetación de la ribera. El procesamiento de las hojas implica medios físicos, químicos y factores biológicos que pueden diferir entre especies vegetales. En esta investigación se evaluó el efecto de la calidad del sustrato sobre el proceso de descomposición y sobre la colonización por invertebrados acuáticos asociados, en un arroyo subtropical. Las hojas de dos especies de árboles nativos, Ficus luschnathiana y Cascarla sylvestris, se incubaron en bolsas de malla gruesa en un arroyo durante 28 días, colectándose muestras semanalmente. Las tasas de descomposición fueron mayores en F. luschnathiana que en C. sylvestris. En los primeros siete días del experimento, F. luschnathiana perdió ~60% de la masa inicial, mientras que C. sylvestris perdió ~30%. Las hojas de C. sylvestris fueron inicialmente más palatables que las de F. luschnathiana, tal vez debido a su menor contenido de lignina y celulosa, y a su más baja relación Lignina:N. Estos compuestos estructurales le conferen rigidez a las hojas, las protegen de los herbívoros y las hacen más resistentes a los detritívoros. Sin embargo, la menor dureza de la hoja de F. luschnathiana facilitó la colonización fúngica, lo cual se refejó, a su vez, en una mayor colonización por invertebrados trituradores y, en general, una mayor pérdida de masa. Los resultados sugieren que la descomposición de la hojarasca fue sensible a la biomasa fúngica y a la dureza de las hojas. Se observó que la colonización microbiana y la disminución de la dureza de las hojas tienen un efecto más fuerte sobre la colonización por invertebrados trituradores y sobre el proceso de descomposición que la calidad química inicial de la hojarasca. La comunidad fúngica es clave en los procesos de degradación en el medio acuático, ya que actúa en la descomposición de la hojarasca, aun en aquella de baja calidad (i.e., F. luschnathiana), promoviendo cambios también en la composición y en la estructura de la comunidad de invertebrados colonizadores.