Abstract Aquatic macrophytes are the main autochthonous component of primary production in the Amazon Basin. Floating meadows of these plants support habitats with highly diverse animal communities. Fishes inhabiting these habitats have been assumed to use a broad range of food items and compose a particular food web. We employed carbon (δ13C) and nitrogen (δ15N) stable isotope analysis to draw the trophic structure of these habitats and to trace the energy flow by its trophic levels. Fishes and other animals from 18 independent macrophyte meadows of a floodplain lake of the Solimões River (Amazonia, Brazil) were analyzed. The food web of macrophyte meadows consists of four trophic levels above autotrophic sources. In general, primary consumers exhibited a broader range of food sources than the upper trophic levels. Some fish species depended on a large number of food sources and at the same time are consumed by several predators. The energy transfer from one trophic level to the next was then mainly accomplished by these species concentrating a high-energy flux and acting as hubs in the food web. The broad range of δ13C values observed indicates that the organisms living in the macrophyte meadows utilize a great diversity of autotrophic sources. Basin communities δC δ C (δ13C δ15N δN N (δ15N 1 Amazonia, Amazonia (Amazonia Brazil analyzed general predators highenergy high

RESUMO O gênero Cichla é bastante diverso, com 16 espécies descritas. Algumas espécies são externamente muito similares e sua discriminação pode ser bastante difícil. Ao mesmo tempo, a discriminação de estoques pesqueiros é essencial para propostas de manejo. Análises morfométricas em otólitos têm sido utilizadas com sucesso para a distinção de espécies e estoques pesqueiros, principalmente em ambientes marinhos. Este estudo avaliou se o formato do otólito sagittal pode ser utilizado para discriminar entre espécies Cichla temensis, C. monoculus e C. orinocensis, bem como dentro de populações de C. temensis em diferentes rios amazônicos. Índices de forma e coeficientes de Fourier foram utilizados para descrever a forma dos otólitos. Dentre as espécies, a morfologia do otólito sagittal do C. temensis mostrou ser totalmente distinta das espécies C. monoculus e C. orinocensis. Enquanto no grupo das populações de C. temensis, os indivíduos dos rios Negro e Jatapú mostraram-se diferentes independente dos métodos utilizados. Esses resultados confirmam a capacidade de diferenciação de espécies e populações através da morfologia dos otólitos. No entanto, são necessárias mais pesquisas para verificar o papel dos efeitos genéticos em comparação aos efeitos ambientais e bióticos para explicar a discriminação observada nos otólitos.

ABSTRACT The genus Cichla is a highly diverse group, with 16 species already described. Externally, some species are very similar and discriminating between them may be very difficult. Nevertheless, discrimination of fish stocks is essential for management purposes. Morphometric analyses of otoliths have been successfully used to distinguish species and fish stocks, especially in marine environments. This study evaluated whether sagittal otolith shape can be used to discriminate among the species Cichla temensis, C. monoculus, and C. orinocensis, as well as within populations of C. temensis in rivers of the Amazon. Shape indices and Fourier coefficients were used to describe the shape of the otoliths. Among the groups of species, the morphology of the sagittal otolith of C. temensis was totally distinct from the species C. monoculus and C. orinocensis. While among populations of C. temensis, individuals from the Negro and Jatapú Rivers were different, regardless of the methods used. These results confirm the ability to differentiate species and populations by using the morphology of otoliths. However, more research is needed to verify the role of genetic versus environmental and biotic effects, and thus be able to explain the discrimination observed in otoliths.

Resumo Semaprochilodus insignis é uma espécie de peixe migratório da Amazônia, movendo-se em cardumes entre rios de água branca e preta. Tem sido classificado como um peixe detritívoro-iliófago. Porém, é possível que a plasticidade trófica de S. insignis seja maior do que se supunha. O objetivo deste estudo foi investigar as contribuições relativas de fontes de energia autotróficas para a dieta do S. insignis nos rios Negro e Solimões e se a espécie muda ontogenticamente a sua dieta. Encontramos variações entre os valores de δ13C e δ15N de S. insignis entre os rios: Negro, o perifíton (84%) foi a principal fonte de energia para adultos, enquanto para os juvenis do rio Solimões, foram predominantemente as plantas terrestres/macrófitas C3 (50%) e fitoplâncton (42%). Essas variações nas assinaturas isotópicas provavelmente estão associadas a movimentos migratórios do S. insignis em diferentes estágios da vida e períodos hidrológicos. Ao invés da classificação alimentar previamente assumida, sugerimos que o S. insignis varie sua dieta ontogeneticamente, os adultos são iliófagos em águas negras, enquanto os jovens são detritívoros em rios de águas brancas. Os resultados demonstram que essa espécie promove ligações complexas entre as cadeias alimentares enfatizando, dessa forma, a importância da conservação das áreas inundadas amazônicas.

Abstract Semaprochilodus insignis is an Amazonian migratory fish species, moving in large shoals between white- and black-water rivers. It has long been classified as a detritivorous fish. However, it is possible that the trophic plasticity of S. insignis could be higher than previously assumed. The objective of this study was to investigate the relative contributions of autotrophic energy sources to the diet of S. insignis in the Negro and Solimões rivers and to determine if the species undergoes an ontogenetic change in the diet. We found variations between the δ13C and δ15N values of S. insignis between the rivers. In the Negro River, periphyton (84%) was the principal energy source for adults, while juveniles in the Solimões River foraged predominantly on terrestrial plants/C3 macrophytes (50%) and phytoplankton (42%). These variations in isotopic signatures are likely associated with migratory movements of S. insignis at different life stages and hydrological periods. Instead of the previously assumed dietary classification, we suggest that S. insignis varies its diet ontogenetically, with adults acting as illiophagous in black-water while the young are detritivorous in white-water rivers. The results show that this species creates complex links between food chains, thus emphasizing the importance of conserving flooded areas.

Con el objetivo de estudiar el efecto de la disponibilidad de recursos sobre la amplitud del nicho trófico en peces de la Amazonia boliviana, comparamos los nichos isotópicos (como sustituto del nicho trófico) de tres especies de peces (Acestrorhynchus altus, Hoplias malabaricus y Pygocentrus nattereri) en un gradiente de disponibilidad de presas. Tomamos muestras de peces en lagos localizados en la llanura de inundación de los ríos Blanco (baja disponibilidad de presas), Iténez (disponibilidad intermedia) y San Martín (disponibilidad alta), analizamos los ratios de isótopos estables (δ13C and δ15N) y estimamos las elipses estándar y amplitud de nicho isotópico (área) empleando el paquete SIBER en R. Acestrorhynchus altus se comportó como un forrajero óptimo, mostrando un nicho isotópico más amplio a menor disponibilidad de presas y un nicho más estrecho en los sitios con disponibilidad intermedia y alta. Pygocentrus nattereri mostró un nicho más amplio a densidades de presa intermedias, y no encontramos efecto de la disponibilidad de presas sobre la amplitud del nicho trófico de H. malabaricus. Esto indica que, para estas dos especies, factores tales como la selectividad de presas, transparencia del agua, tamaño de la boca y hábitos alimenticios, podrían ser más importantes que la disponibilidad de presas para determinar la amplitud del nicho trófico.

To study the effect of resource availability on the trophic niche width of fishes in the Bolivian Amazon basin, we compared isotopic niche widths (as a surrogate for trophic niche) of three fish species (Acestrorhynchus altus, Hoplias malabaricus and Pygocentrus nattereri) in a gradient of prey availability. We sampled fishes in lakes located in the floodplain of the rivers Blanco (low prey availability), Iténez (intermediate prey availability) and San Martín (high prey availability), analyzed them for ratios of stable isotopes (δ13C and δ15N), and estimated the standard ellipse and isotopic niche width using the SIBER package in R. Acestrorhynchus altus behaved as an optimal forager, showing wider isotopic niche where prey were less abundant, and less wide isotopic niche in rivers with intermedíate and high prey availability. P. nattereri showed wider trophic niche at intermedíate prey densities and H. malabaricus showed no effect of prey availability on isotopic niche width. This indicates that, for these species, factors such as prey selectivity, water transparency, mouth size or feeding habits, may be more important than prey availability in determining trophic niche width.

ResumenLos gradientes altitudinales representan un sistema apropiado para evaluar si existe una relación general entre los patrones de riqueza, variables ambientales, y los procesos ecológicos que determinan cuantas especies co-ocurren en un área dada.En cuanto a los peces de agua dulce en arroyos de montaña la relación prevalente es un decrecimiento monotónico en riqueza de especies con la altura. El objetivo de este estudio es evaluar cuatro hipótesis que pueden explicar la relación negativa entre la riqueza local de especies de peces y la altura, 1) hipótesis de la disminución de la energía disponible 2) hipótesis del aumento del rigor climático 3) hipótesis de la diversidad de hábitats e 4) hipótesis de la severidad de las condiciones físicas del hábitat que se presentan río arriba, relacionada a las dificultades para colonizar las partes altas de los ríos. Se recolectaron peces y macro-invertebrados en 83 sitios entre 200 a 4 000 m en dos cuencas en la Amazonía boliviana. La primera hipótesis fue evaluada mediante regresiones entre la densidad de macro-invertebrados, los peces invertívoros y la altura. Para evaluar la segunda y tercera hipótesis se realizó un análisis de regresión múltiple (GLM), entre la riqueza y varios factores locales y regionales. Se analizó también la disimilitud de los ensamblajes de peces sobre el gradiente altitudinal utilizando los índices βsim y βnes. Se encontró que la riqueza de peces disminuye de manera lineal con el aumento de la altura. La densidad de macroinvertebrados aumenta con la altura, contrariamente a la riqueza de peces invertívoros, sugiriendo que la energía no constituye un factor limitante para la colonización de las especies de peces en zonas altas. El GLM explicó un 86 % de la variación total de la riqueza, con un aporte significativo de la temperatura del agua, las pendientes máximas en el río principal y el ancho de río. El βsim presentó valores altos en niveles de altura bajos, mostrando mayor recambio de especies entre los sitios cercanos a la fuente de colonización. Inversamente, el βnes mostró valores mayores en las partes altas, correspondientes a cambios por pérdida de especies. Estos resultados sugieren que el rigor climático junto con la severidad física del hábitat crean barreras a la colonización y explican los cambios de riqueza en el gradiente altitudinal.

AbstractAltitudinal gradients represent an appropriate system to assess whether there is a relationship between richness patterns, environmental variables, and the ecological processes that determine the species type and number inhabiting a given area. In mountain streams freshwater fishes, the most prevalent relationship is a monotonic decrease in species richness with elevation. The objective of this study was to evaluate four hypotheses that can explain the negative relationship between local fish species richness and altitude, 1) the hypothesis of decreasing energy availability, 2) the hypothesis of increasing climate severity, 3) the hypothesis of habitat diversity, and 4) the hypothesis of isolation by physical severity of the environment. Fish and macro-invertebrates were collected following standard methods from 83 sites (between 200-4 000 meters) of two river basins in the Bolivian Amazon. The first hypothesis was tested by analyzing relationships between the density of macro-invertebrates, the richness of invertivorous fish species and altitude; while the second and third hypotheses were assessed by a multiple regression analysis (GLM) between fish species richness and several local and regional factors. Besides, assemblage dissimilarity between sites along the altitudinal gradient was analyzed using βsim and βness indices. Fish richness decreases linearly with increasing altitude. The density of macro-invertebrates tends to increase at higher altitudes, contrary to invertivorous fish species richness, suggesting that energy availability is not a limiting factor for fish species colonization. The GLM explained 86 % of the variation in fish species richness, with a significant contribution of water temperature, maximum slope in the river mainstem, and stream width. There is a higher species turnover (βsim) between sites at low elevation. Inversely, βness shows higher values in the upper parts, corresponding to change in assemblages mainly due to species loss. Taken together, these results suggest that climatic and physical severities create strong barriers to colonization, further explaining the decrease in fish richness along the altitudinal gradient.