Abstract
The drift movement consists of the displacement of the organisms inside the water column which allows its passive locomotion. This movement will result in a variation of the communities of organisms along the river, generating spatial patterns. Based on this, we tested the hypotheses a) the drift of individuals in an upstream-downstream direction creates a nestedness pattern, when the upstream is a subset of downstream communities of aquatic insects; b) there will be an increase in the number of individuals and genera as we approach the most downstream point. The present study was carried out in seven sampling points distributed along the Xingu River. The sampling occurred at night in the central area of the river. The number of genera along the river remained constant, and the nestedness distribution of the communities in the upstream-downstream gradient was not observed. Based on the results, it is possible to visualize a turnover of genera in the longitudinal gradient of the river, but with an accumulation of genera in the downstream region. Organisms that are transported by the flow of the water current respond to the characteristics of the body of water by adapting to the type of environment in which they are found.
Keywords
Dispersion; Aquatic Insects; Amazonian River; Upstream-downstream movement
Resumo
O movimento de deriva consiste no desprendimento dos organismos dentro da coluna de água, o que permite a sua locomoção passiva. Este movimento resultará numa variação das comunidades de organismos ao longo do rio, gerando padrões espaciais. Com base nisto, testamos as hipóteses a) o movimento de indivíduos em direção montante-jusante criará um padrão aninhado, no qual as comunidades de insetos aquáticos a montante são um subconjunto das comunidades a jusante; b) haverá um aumento no número de indivíduos e gêneros à medida que nos aproximamos do ponto mais a jusante. O presente estudo foi realizado em sete pontos de amostragem distribuídos ao longo do rio Xingu. A amostragem ocorreu durante a noite no canal central do rio. O número de gêneros ao longo do rio se manteve constante, e não observamos uma distribuição de aninhamento das comunidades no gradiente ascendente e descendente do rio. Com base nos resultados, é possível visualizar uma substituição dos gêneros no gradiente longitudinal do rio, porém ocorrendo um acúmulo de gêneros na região mais a jusante. Os organismos que são transportados pelo fluxo da corrente de água respondem as características do corpo de água adaptando-se ao tipo de ambiente em que se encontram.
Palavras-chave
Dispersão; Insetos Aquáticos; Rio Amazônico; Movimento montante-jusante
Introduction
Water flow plays a major role in the dynamics of lotic environments (e.g., rivers and streams) and is related to the stability of biological populations (Poff & Ward 1991POFF, N.L. & WARD, J. V. 1991. Drift Responses of Benthic Invertebrates to Experimental Streamflow Variation in a Hydrologically Stable Stream. Can. J. Fish. Aquat. Sci. 48(10):1926–1936.). The movement of water promotes the drift of organisms, which consists of their transport using the water flow. The drift behavior is related to several factors, such as current velocity, water chemistry, period of the year and photoperiod (Fierro et al. 2015FIERRO, P., BERTRAN, C., MERCADO, M., PENA CORTES, F., TAPIA, J., HAUENSTEIN, E., CAPUTO, L. & VARGAS CHACOFF, L. 2015. Landscape composition as a determinant of diversity and functional feeding groups of aquatic macroinvertebrates in southern rivers of the Araucania, Chile. Lat. Am. J. Aquat. Res. 43(1):186–200.). The drift can be classified as active, when the organism is cast into the water column in order to escape from predation, competition, or seek food; or passive when the organism is involuntarily thrown into the water column (Brittain & Eikeland 1988BRITTAIN, J.E. & EIKELAND, T.J. 1988. Invertebrate drift - A review. Hydrobiologia 166(1):77–93., Poff & Ward 1991POFF, N.L. & WARD, J. V. 1991. Drift Responses of Benthic Invertebrates to Experimental Streamflow Variation in a Hydrologically Stable Stream. Can. J. Fish. Aquat. Sci. 48(10):1926–1936., Castro et al. 2013aCASTRO, D., HUGHES, R. & CALLISTO, M. 2013a. Influence of peak flow changes on the macroinvertebrate drift downstream of a Brazilian hydroelectric dam. Brazilian J. Biol. 73(4):775–782.).
The study of the drift movement is fundamental to understand the transport of these organisms through the water flow, to understand the process of colonization and recolonization of habitats, as well as to identify the functional ecosystem role of different species. The drift movement of aquatic insects can be responsible for the stability and structure of the communities existing in each environment through the process of repopulating localities (Vellend 2010VELLEND, M. 2010. Conceptual synthesis in community ecology. Q. Rev. Biol. 85183–206.). There is this dependence because the distribution of the species is also linked to the environmental variables and to the physiological tolerance of each organism (Godoy, Queiroz, et al. 2022GODOY, B.S., QUEIROZ, L.L., SIMIÃO-FERREIRA, J., LODI, S., CAMARGOS, L.M. & OLIVEIRA, L.G. 2022. The effect of spatial scale on the detection of environmental drivers on aquatic insect communities in pristine and altered streams of the Brazilian Cerrado. Int. J. Trop. Insect Sci. 42(3):2173–2182., Vellend 2010VELLEND, M. 2010. Conceptual synthesis in community ecology. Q. Rev. Biol. 85183–206.). Thus, studies on drift movement are being used to understand the distribution of organisms in order to determine the connection between localities (Poff & Ward 1991POFF, N.L. & WARD, J. V. 1991. Drift Responses of Benthic Invertebrates to Experimental Streamflow Variation in a Hydrologically Stable Stream. Can. J. Fish. Aquat. Sci. 48(10):1926–1936., Anholt 1995ANHOLT, B.R. 1995. Density Dependence Resolves the Stream Drift Paradox. Ecology 76(7):2235–2239., Covich 2006COVICH, A.P. 2006. Dispersal - limited biodiversity of tropical insular streams. Polish J. Ecol. 54(4):523–547.).
The flow of water into a river is unidirectional, so it is expected in an upstream-downstream direction for the transport of organisms carried by the stream. This will generate a pattern of species distribution, of species addition in the upstream-downstream gradient. The pattern generated may also be of the nestedness type, in which the upstream communities are a subset of those found downstream (Covich 2006COVICH, A.P. 2006. Dispersal - limited biodiversity of tropical insular streams. Polish J. Ecol. 54(4):523–547., Almeida-Neto et al. 2008ALMEIDA-NETO, M., GUIMARÃES, P., GUIMARÃES, P.R., LOYOLA, R.D. & ULRICH, W. 2008. A consistent metric for nestedness analysis in ecological systems: reconciling concept and measurement. Oikos 117(8):1227–1239.). The concept of nestedness was created to explain the communities of island colonized by species from mainland, wherein the better disperses colonize the great portion of islands, independent of the distance (Darlington 1966DARLINGTON, P.J. 1966. Zoogeography: the geographical distribution of animals. 1 ed. Wiley & Sons/Museum of Comparative Zoology/Harvard University, Cam.). In other hand, the poor disperses occur only in the more connected island, creating a nested pattern of species occurrences, based in the species incidence. The knowledge about nestedness distribution is important to understand the patterns of community composition in ecosystems and based on this knowledge create strategies for preservation and conservation of the environment (Ulrich 2009ULRICH, W. 2009. Nestedness analysis as a tool to identify ecological gradients. Ecol. Quest. 11(1):). However, most of the studies on drift movement are concentrated in streams, resulting in a knowledge gap in relation to large rivers that present a differentiated dynamic, with a more intense water flow and variation between environments.
The aquatic insects are strongly influenced by the water dynamics of the water bodies and, in order to establish themselves in the environment, use adapted mechanisms to the conditions to which they are submitted (Mazzucco et al. 2015MAZZUCCO, R., VAN NGUYEN, T., KIM, D.-H., CHON, T.-S. & DIECKMANN, U. 2015. Adaptation of aquatic insects to the current flow in streams. Ecol. Modell. 309–310143–152.). In addition to the water flow, these organisms suffer interference from the substrate type, and rocky substrates have a differentiated community when compared to sandy environments (Bispo et al. 2004BISPO, P., OLIVEIRA, L., CRISCI-BISPO, V. & SOUSA, K. 2004. Environmental Factors Influencing Distribution and Abundance of Trichopteran Larvae in Central Brazilian Mountain Streams. Stud. Neotrop. Fauna Environ. 39(3):233–237.). The availability of food and predation will also be limiting factors and may define the population density of each locality (Ciborowski 1983CIBOROWSKI, J.J.H. 1983. Influence of current velocity, density, and detritus on drift of two mayfly species (Ephemeroptera). Can. J. Zool. 61(1):119–125., Hay et al. 2008HAY, C.T., CROSS, P.C. & FUNSTON, P.J. 2008. Trade-offs of predation and foraging explain sexual segregation in African buffalo. J. Anim. Ecol. 77(5):850–858., Godoy et al. 2016GODOY, B.S., QUEIROZ, L.L., LODI, S., NASCIMENTO DE JESUS, J.D. & OLIVEIRA, L.G. 2016. Successional colonization of temporary streams: An experimental approach using aquatic insects. Acta Oecologica 7743–49.). Environments with high population density show an increase in biotic interactions, causing the escape of individuals who are forced into the water column. The active dispersal movement occurs more intensely during the night period, where the lack of luminosity provides a protection against the predators, guaranteeing a greater success during the displacement in the water column (Bishop 1969BISHOP, J.E. 1969. Light Control of Aquatic Insect Activity and Drift. Ecology 50(3):371–380., Koetsier 2005KOETSIER, P. 2005. The Effects of Disturbance Time Interval on Algal Biomass in a Small Idaho Stream. Northwest Sci. 79(4):211–217.). There are many studies with the orders Ephemeroptera, Plecoptera and Trichoptera (EPT) about their distribution and life cycle (Godoy, Valente‐Neto, et al. 2022GODOY, B.S., VALENTE‐NETO, F., QUEIROZ, L.L., HOLANDA, L.F.R., ROQUE, F.O., LODI, S. & OLIVEIRA, L.G. 2022. Structuring functional groups of aquatic insects along the resistance/resilience axis when facing water flow changes. Ecol. Evol. 12(3):., Godoy, Queiroz, et al. 2022GODOY, B.S., QUEIROZ, L.L., SIMIÃO-FERREIRA, J., LODI, S., CAMARGOS, L.M. & OLIVEIRA, L.G. 2022. The effect of spatial scale on the detection of environmental drivers on aquatic insect communities in pristine and altered streams of the Brazilian Cerrado. Int. J. Trop. Insect Sci. 42(3):2173–2182., Merritt et al. 2008MERRITT, R.W., CUMMINS, K.W. & BERG, M.B. 2008. An introduction to the aquatic insects of North America. 4 ed. Kendall / Hunt Publishing Company, Dubuque., Sarremejane et al. 2020SARREMEJANE, R., CID, N., DATRY, T., STUBBINGTON, R., ALP, M., CAÑEDO-ARGÜELLES, M., CORDERO-RIVERA, A., CSABAI, Z., GUTIÉRREZ-CÁNOVAS, C., HEINO, J., FORCELLINI, M., MILLÁN, A., PAILLEX, A., PAŘIL, P., POLÁŠEK, M., FIGUEROA, J.M.T. de, USSEGLIO-POLATERA, P., ZAMORA-MUÑOZ, C. & BONADA, N. 2020. DISPERSE: A trait database to assess the dispersal potential of aquatic macroinvertebrates. Sci. Data 7386.). In addition, they are orders with high environmental sensitivity, wide distribution within the lotic environments, high abundance and each order presents high richness and complexity (Godoy et al. 2019GODOY, B.S., FARIA, A.P.J., JUEN, L., SARA, L. & OLIVEIRA, L.G. 2019. Taxonomic sufficiency and effects of environmental and spatial drivers on aquatic insect community. Ecol. Indic. 107105624.), which allows its use as model organisms for studies of the drift process.
The Xingu River is characterized as a large river, presenting along its length a high environmental heterogeneity, characterized as a river of high complexity. Its landscapes are composed by waterfalls and rapids, where in these localities the speed of the current appears very varied. In addition to this variation of the current, there will be a variation of the biological communities within its course, resulting in a distribution of varied species within each environment. The lack of studies in large-scale rivers undermines understanding of ecosystem dynamics and how changes in water flow alter the structure of aquatic insect communities. Thus, in this study we tested the following hypotheses: 1) there will be a nesteness distribution of the genera Ephemeroptera and Trichoptera in an upstream-downstream gradient along the Xingu River; 2) there will be an increase in the abundance and richness of genera as we approach the most downstream point on the Xingu River, and c) the community of aquatic insects moves upstream.
Material and Methods
1. Study area
The study was carried out on the Xingu River (02°51′33.1″S and 52°19′28″W), near the city of Altamira, Pará, during the flood period, April 2015 (Figure 1). The Xingu River belongs to the Amazon River Basin, located on the right side of the river. With an extension of 1500 km from its source in the Brazilian Central Plateau until its mouth in the Amazon River, it drains an area of 540 km2. The pH ranges from 5.5 to 7.0 with a mean conductivity of 30 μS/cm–1, as well as high concentrations of oxygen resulting from the large volume of water (Sioli 1957SIOLI, H. 1957. Valores de pH de águas Amazônicas. Bol. do Mus. Para. Emilio Goeldi 11–35., Salomão et al. 2007SALOMÃO, R.P., VIEIRA, I.C.G., SUEMITSU, C., ROSA, N. de A., ALMEIDA, S.S. de, AMARAL, D.D. do & MENEZES, M.P.M. de. 2007. As florestas de Belo Monte na grande curva do rio Xingu, Amazônia Oriental. Bol. do Mus. Para. Emílio Goeldi, Ciências Nat. 2(3):57–153.).
Sampling points of aquatic insects on the Xingu River in the period of April 2015. The sample units are symbolized by the P code.
The average flow rate during the flood period varies from 8,000 to 10,000 m3/s and in the dry period the average is 2000 m3/s (Norte-Energia 2016NORTE-ENERGIA. 2016. UHE Belo Monte.). The flood period occurs between December and April and the dry season occurs between July and November. Because it is located near the equator, the Xingu River basin presents a warm climate and according to Köppen classification the climate is tropical and predominantly humid (Am, Sheffield et al. 2006SHEFFIELD, J., GOTETI, G. & WOOD, E.F. 2006. Development of a 50-Year High-Resolution Global Dataset of Meteorological Forcings for Land Surface Modeling. J. Clim. 19(13):3088–3111.). The mean annual temperature in the Altamira-PA region is 27°C, the rainy season starts in November and the dry season in July.
2. Sampling
The sampling occurred in seven locations on the Xingu River during the night period. We sampled two times in the central region of the river in each location. However, for our study we jointed the two samples. At each sampling point we measured abiotic variables (pH, OD, conductivity, temperature and current velocity). We used a plankton net with a 50 cm diameter ring and 1.5 m in length, with a mesh opening of 300 μm (Bialetzki et al. 1999BIALETZKI, A., SANCHES, P.V., CAVICCHIOLI, M., BAUMGARTNER, G., RIBEIRO, R.P. & NAKATANI, K. 1999. Drift of ichthyoplankton in two channels of the Paraná River, between Paraná and Mato Grosso do Sul states, Brazil. Brazilian Arch. Biol. Technol. 42(1):.). We attached a weight to the net, aiming at its balance in the water column. A flowmeter was attached to the net to determine the amount of filtered water and at the end of the net a collecting cup was added. We performed the samplings with the aid of a motorized canoe, which during the collection was kept on with low acceleration, with the bow upstream of the river. The net was positioned against the current, at an average depth of 2 meters, for a period of 10 minutes, adapting methodologies previously used in works with aquatic insects (Waters 1972WATERS, T.F. 1972. The Drift of Stream Insects. Annu. Rev. Entomol. 17(1):253–272., Castro et al. 2013aCASTRO, D., HUGHES, R. & CALLISTO, M. 2013a. Influence of peak flow changes on the macroinvertebrate drift downstream of a Brazilian hydroelectric dam. Brazilian J. Biol. 73(4):775–782., bCASTRO, D.M.P., HUGHES, R.M. & CALLISTO, M. 2013b. Effects of flow fluctuations on the daily and seasonal drift of invertebrates in a tropical river. Ann. Limnol. Int. J. Limnol. 49(3):169–177.). The sampled individuals were preserved in 70% ethanol and identified to the genus level (Wiggins 1977WIGGINS, G.B. 1977. Larvae of the North American caddisfly genera (Trichoptera). University of Toronto Press, Toronto., Domingues & Fernandez 2001DOMINGUES, E. & FERNANDEZ, H. 2001. Guia para la determinación de los artrópodos bentónicos sudamericanos. Universidad Nacional de Tucumán, Tucumán., Pes et al. 2005PES, A.M.O., HAMADA, N. & NESSIMIAN, J.L. 2005. Chaves de identificação de larvas para famílias e gêneros de Trichoptera (Insecta) da Amazônia Central, Brasil. Rev. Bras. Entomol. 49(2):181–204., Oliveira 2006OLIVEIRA, L.G. 2006. Trichoptera. In Insetos imaturos - Metamorfose e identificação (C. Costa, S. Ide, & C. Simonka, eds) Holos, Ribeirão Preto.).
3. Data analysis
To verify the relationship between the number of genera and the abundance of individuals with the upstream-downstream gradient of the points sampled, a linear regression was performed, using the ordering of the samples in this gradient as a predictor variable. We used NODF (Nestedness metric based on Overlapping and Decreasing Fill) (Almeida-Neto et al. 2008ALMEIDA-NETO, M., GUIMARÃES, P., GUIMARÃES, P.R., LOYOLA, R.D. & ULRICH, W. 2008. A consistent metric for nestedness analysis in ecological systems: reconciling concept and measurement. Oikos 117(8):1227–1239.), to observe a nestedness pattern in the upstream-downstream gradient for the aquatic insect community within the river course. This metric works in a range between 0-100, with 100 representing a perfectly nesteness set. The data are organized in an array of rows and columns, where the columns are the genera and in the rows are the points sampled, the NODF is calculated in pairs of subsequent rows and columns, if the previous row has lesser or equal number of genera than the after, the value of the NODF will be zero. However, if the previous line has higher number of genera than the posterior one, the index uses the common occurrence in both lines to generated the value of the NODF. This calculation is performed for both rows and columns and at the end the overall mean will be achieved resulting in the general NODF (Milesi & Melo 2014MILESI, S.V. & MELO, A.S. 2014. Conditional effects of aquatic insects of small tributaries on mainstream assemblages: position within drainage network matters Y. Chen, ed. Can. J. Fish. Aquat. Sci. 71(1):1–9., Pinha et al. 2016PINHA, G.D., PETSCH, D.K., RAGONHA, F.H., GUGLIELMETTI, R., BILIA, C.G., TRAMONTE, R.P. & TAKEDA, A.M. 2016. Benthic invertebrates nestedness in flood and drought periods in a Neotropical floodplain: looking for the richest environments. Acta Limnol. Bras. 28.). In order to calculate the NODF in our study, we maintained the order of the sampling sites by fixing the points in the upstream-downstream direction, generating the real distribution of the genera of aquatic insects in the studied gradient. We performed the T test to compare the observed NODF values with the estimated distribution in a null model of 1000 iterations for the NODF values.
The dispersion among the communities of the points sampled was determined by the dispersion coefficient of the biogeographic direction, using the DD3 index (Legendre & Legendre 1984LEGENDRE, P. & LEGENDRE, V. 1984. Postglacial Dispersal of Freshwater Fishes in the Québec Peninsula. Can. J. Fish. Aquat. Sci. 41(12):1781–1802.). This index determines in what direction the movement of species is occurring between communities of locations that are geographically connected (Borcard et al. 1995BORCARD, D., GEIGER, W. & MATTHEY, W. 1995. Oribatid mite assemblages in a contact zone between a peat-bog and a meadow in the Swiss Jura (Acari, Oribatei): inflence of landscape structures and historical process. Pedobiologia (Jena). 39318–330., Legendre & Legendre 1984LEGENDRE, P. & LEGENDRE, V. 1984. Postglacial Dispersal of Freshwater Fishes in the Québec Peninsula. Can. J. Fish. Aquat. Sci. 41(12):1781–1802.). For the analysis, the Vegan package was used (Oksanen et al. 2013OKSANEN, J., BLANCHET, F.G., KINDT, R., LEGENDRE, P., MINCHIN, P.R., O’HARA, R.B., SIMPSON, G.L., SOLYMOS, P., STEVENS, M.H.H. & WAGNER, H. 2013. Package ‘vegan.’ R Packag. ver. 2.0–8 254.) available in the R program (R Development Core Team 2020R DEVELOPMENT CORE TEAM. 2020. R Development Core Team, R: a language and environment for statistical computing.).
Results
We collected 1760 individuals in total, divided into 13 families and 34 genera. The order Ephemeroptera consisted of 1614 individuals divided into 6 families and 21 genera. The order Trichoptera presented 146 individuals distributed in 7 families and 13 genera. The most abundant genera were Lachlania, Camelobaetidius, Hydrosmilodon and Cloeodes (Table 1). The families that obtained the highest representation were Baetidae (31.2%), Oligoneuriidae (29.01%) and Leptophlebiidae (22.26%). The values of physical and chemical variables showed low variability between the locations (Table 2).
Aquatic insects’ genera, abundance and occurrence in samples collected in the channel of Xingu River, during the flood period (April/2015).
The number of genera between the sampling points did not differ significantly, except for the last sampling point located downstream, where there was a reduction in the number of genera (F1,5 = 1.99, p = 0.28). However, this reduction should not be interpreted as an indication of decay, since the pattern is not clear, and more collections are needed before we can even visualize this distribution. The same pattern was observed for the abundance of individuals following the downstream gradient (F1,5 = 0.01, p = 0.90).
The distribution of the genera showed a turnover pattern (NODF observed: 28.88; NODF estimated: 58.51; T = 7.97, p < 0.01), wherein a genera substitution and each sampling point there is a differentiated community. The genera Needhamella, Lachlania, Hydrosmilodon, Cloeodes, Campsurus, Camelobaetis, Tricorythopsis were the most expressive in the abundance being present at all sampling points, whereas the genera Anacroneuria, Austrotinodes, Chimarra, Hagenulopsis, Caenis, Cryptonympha, Nectopsyche, Hydroptila, Neotrichia, Tikuna, Ulmeritoides and Tricorythodes occurred at only one sampling point.
The result of the dispersion coefficient (DD3 index) indicated a possible existence of a dispersion pattern in the downstream direction of the river, where the organisms move following the current flow. Individuals were found to carry out a dispersion movement directed towards the last sampling point located further downstream (Figure 2). The positive values in the table indicate that the displacement is occurring in a downstream direction while negative values indicate that a reverse movement occurs (Table 3).
Dispersion between communities of the Xingu River in a downstream gradient. At the top of the table are the values for index DD3, while at the bottom are the associated p values. Bold values indicate p < 0.05
Discussion
The dispersion movement within aquatic insect communities is responsible for the interaction between communities from different localities, resulting in the colonization of environments and increasing local diversity. The physical and chemical characteristics of each locality will generate different organizations of aquatic insects, where the most adapted to a certain condition will be able to establish themselves (Rodrigues-Filho et al. 2015RODRIGUES-FILHO, J., ABE, D., GATTI-JUNIOR, P., MEDEIROS, G., DEGANI, R., BLANCO, F., FARIA, C., CAMPANELLI, L., SOARES, F., SIDAGIS-GALLI, C., TEIXEIRA-SILVA, V., TUNDISI, J., MATSMURA-TUNDISI, T. & TUNDISI, J. 2015. Spatial patterns of water quality in Xingu River Basin (Amazonia) prior to the Belo Monte dam impoundment. Brazilian J. Biol. 75(3 suppl 1):34–46.). The aquatic insect communities that perform their locomotion through the water column in the Xingu River did not demonstrate a nestedness distribution, refuting our first hypothesis. In order to occur a nestedness distribution, the community of aquatic insects found downstream should be a subset of the community found downstream, however we observed a species substitution for each sampling point, a pattern normally observed when the environment is relevant to the community structure (Heino 2009HEINO, J. 2009. Biodiversity of Aquatic Insects: Spatial Gradients and Environmental Correlates of Assemblage-Level Measures at Large Scales. Freshw. Rev. 2(1):1–29.).
The results obtained in this study showed no variation in the number of insect genera along the course of the river, resulting in a uniformity in the community of aquatic insects. The environmental variables don`t showed a relevant variation between the sample points, and the marginal vegetation surround the river is well preserved. The marginal vegetation is a great driver in the chemical conditions of the water (Salomão et al. 2007SALOMÃO, R.P., VIEIRA, I.C.G., SUEMITSU, C., ROSA, N. de A., ALMEIDA, S.S. de, AMARAL, D.D. do & MENEZES, M.P.M. de. 2007. As florestas de Belo Monte na grande curva do rio Xingu, Amazônia Oriental. Bol. do Mus. Para. Emílio Goeldi, Ciências Nat. 2(3):57–153., Sawakuchi et al. 2015SAWAKUCHI, A.O., HARTMANN, G.A., SAWAKUCHI, H.O., PUPIM, F.N., BERTASSOLI, D.J., PARRA, M., ANTINAO, J.L., SOUSA, L.M., SABAJ PÉREZ, M.H., OLIVEIRA, P.E., SANTOS, R.A., SAVIAN, J.F., GROHMANN, C.H., MEDEIROS, V.B., MCGLUE, M.M., BICUDO, D.C. & FAUSTINO, S.B. 2015. The Volta Grande do Xingu: reconstruction of past environments and forecasting of future scenarios of a unique Amazonian fluvial landscape. Sci. Drill. 2021–32.), and the lack of major alterations in the course of sampling points and due to this uniformity in the environment, may be results in the stability of aquatic insect communities. As water level rises, there will be a greater interconnection between the environments facilitating the movement of dispersion between the environments (Wilson & McTammany 2016WILSON, M.J. & MCTAMMANY, M.E. 2016. Spatial scale and dispersal influence metacommunity dynamics of benthic invertebrates in a large river. Freshw. Sci. 35(2):738–747., Barbosa et al. 2015BARBOSA, T., BENONE, N., BEGOT, T., GONÇALVES, A., SOUSA, L., GIARRIZZO, T., JUEN, L. & MONTAG, L. 2015. Effect of waterfalls and the flood pulse on the structure of fish assemblages of the middle Xingu River in the eastern Amazon basin. Brazilian J. Biol. 75(3 suppl 1):78–94.).
However, the variability of environmental conditions in the Xingu River is related to the seasonality (Salomão et al. 2007SALOMÃO, R.P., VIEIRA, I.C.G., SUEMITSU, C., ROSA, N. de A., ALMEIDA, S.S. de, AMARAL, D.D. do & MENEZES, M.P.M. de. 2007. As florestas de Belo Monte na grande curva do rio Xingu, Amazônia Oriental. Bol. do Mus. Para. Emílio Goeldi, Ciências Nat. 2(3):57–153., Sawakuchi et al. 2015SAWAKUCHI, A.O., HARTMANN, G.A., SAWAKUCHI, H.O., PUPIM, F.N., BERTASSOLI, D.J., PARRA, M., ANTINAO, J.L., SOUSA, L.M., SABAJ PÉREZ, M.H., OLIVEIRA, P.E., SANTOS, R.A., SAVIAN, J.F., GROHMANN, C.H., MEDEIROS, V.B., MCGLUE, M.M., BICUDO, D.C. & FAUSTINO, S.B. 2015. The Volta Grande do Xingu: reconstruction of past environments and forecasting of future scenarios of a unique Amazonian fluvial landscape. Sci. Drill. 2021–32.). During the rainy season the river showed an increase of environmental heterogeneity, because large portions of the bottom of the river (included big stones and mud stretch) were exposed out of the water. In other hand, during the period of flood occurs a uniformity of the environments, because the water covers great portion of the margins, and in the period of drought the environment becomes more heterogeneous with the emergence of rocky outcrops that will make the flow of water be varied (Sawakuchi et al. 2015SAWAKUCHI, A.O., HARTMANN, G.A., SAWAKUCHI, H.O., PUPIM, F.N., BERTASSOLI, D.J., PARRA, M., ANTINAO, J.L., SOUSA, L.M., SABAJ PÉREZ, M.H., OLIVEIRA, P.E., SANTOS, R.A., SAVIAN, J.F., GROHMANN, C.H., MEDEIROS, V.B., MCGLUE, M.M., BICUDO, D.C. & FAUSTINO, S.B. 2015. The Volta Grande do Xingu: reconstruction of past environments and forecasting of future scenarios of a unique Amazonian fluvial landscape. Sci. Drill. 2021–32.). It was found in this study that during the flood period the structure of the aquatic insect community does not show large changes between the sampling points, and studies are needed to compare how insect community structuring occurs during the dry season.
Our results indicated the last downstream point as the destination of the genera. All genera observed at the most upstream points are scattering to this location, generating a drift pattern of genera that tend to follow the flow of the river. The dispersion movement is necessary for the colonization of new areas, in view of the fact that communities are interlinked to the dispersion movement and aim to establish the equilibrium of populations in the environment, keeping population sizes in line with environmental support capacity (Waters 1972WATERS, T.F. 1972. The Drift of Stream Insects. Annu. Rev. Entomol. 17(1):253–272., Mazzucco et al. 2015MAZZUCCO, R., VAN NGUYEN, T., KIM, D.-H., CHON, T.-S. & DIECKMANN, U. 2015. Adaptation of aquatic insects to the current flow in streams. Ecol. Modell. 309–310143–152.). However, that statement is need to be tested using other methods to verify the locomotion of individuals, like mark-recapture methods. In addition, we need caution to interpret the result of our study, because the reduced number of sample locations (seven) may be representing a relevant, but limited, source of information about this aquatic community.
The presence of aquatic insects in the water column has ecological relevance, such as food for larger organisms such as fish, processing of organic matter and provides energy within the trophic web. The location of the occurrence of aquatic insects in the regions of the river column may be related to the life strategies of the animals, where they can counterbalance the food encounter with the dispersion and leakage capacity provided by the water flow, in the water column due to poor fixation on the substrate. Understanding how the dispersion movement occurs is necessary so that we can understand patterns of distribution, richness and interaction between species from different localities (Junior & Suarez 2015JUNIOR, W. & SUAREZ, Y. 2015. Metacomunidades em riachos: Uma abordagem cienciométrica. Biodiversidade 1432–42.).
The EPT orders have a wide global distribution and there are many studies with this group about its ecology and bionomics characteristics (Marques et al. 1999MARQUES, M.G.S.M., FERREIRA, R.L. & BARBOSA, F.A.R. 1999. A comunidade de macroinvertebrados aquáticos e características limnológicas das lagoas Carioca e da Barra, Parque Estadual do Rio Doce, MG. Rev. Bras. Biol. 59(2):203–210., Baumgartner et al. 2004BAUMGARTNER, G., NAKATANI, K., GOMES, L., BIALETZKI, A. & SANCHES, P. 2004. Identification of spawning sites and natural nurseries of fishes in the upper Paraná River, Brazil. Environ. Biol. Fishes 71(2):115–125., Galdean et al. 2001GALDEAN, N., CALLISTA, M. & BARBOSA, F.A.R. 2001. Biodiversity assessment of benthic macroinvertebrates in altitudinal lotic ecosystems of Serra do Cipó (MG, Brazil). Rev. Bras. Biol. 61(2):239–248., Gualdoni & Oberto 2012GUALDONI, C.M. & OBERTO, A.M. 2012. Estructura de la comunidad de macroinvertebrados del arroyo Achiras (Córdoba, Argentina): análisis previo a la construcción de una presa. Iheringia. Série Zool. 102(2):177–186., Godoy, Valente‐Neto, et al. 2022GODOY, B.S., VALENTE‐NETO, F., QUEIROZ, L.L., HOLANDA, L.F.R., ROQUE, F.O., LODI, S. & OLIVEIRA, L.G. 2022. Structuring functional groups of aquatic insects along the resistance/resilience axis when facing water flow changes. Ecol. Evol. 12(3):., Godoy, Queiroz, et al. 2022GODOY, B.S., QUEIROZ, L.L., SIMIÃO-FERREIRA, J., LODI, S., CAMARGOS, L.M. & OLIVEIRA, L.G. 2022. The effect of spatial scale on the detection of environmental drivers on aquatic insect communities in pristine and altered streams of the Brazilian Cerrado. Int. J. Trop. Insect Sci. 42(3):2173–2182.). We can better understand the dynamics of dispersion and drift when we observe the characteristics of each order separately. The order Ephemeroptera is strongly influenced by the organic matter present in association with this type of material (Hamada et al. 2019HAMADA, N., NESSIMIAN, J. & QUERINO, R. 2019. Insetos Aquáticos na Amazônia brasileira: taxonomia, biologia e ecologia.). The samplings were carried out during the flood period when there is a large quantity of organic matter. The high level of the river transports the organic matter present in the marginal region to the channel resulting in an increase in the passive drift of these organisms, besides this factor, we have the high-water flow that exerts strong pressure for the detachment of these organisms from the substrate. Due to these conditions, the order Ephemeroptera was the most abundant and diverse in our samples (Bauernfeind & Moog 2000BAUERNFEIND, E. & MOOG, O. 2000. Mayflies (Insecta: Ephemeroptera) and the assessment of ecological integrity: A methodological approach. Hydrobiologia 422/42371–83., Barbosa et al. 2015BARBOSA, T., BENONE, N., BEGOT, T., GONÇALVES, A., SOUSA, L., GIARRIZZO, T., JUEN, L. & MONTAG, L. 2015. Effect of waterfalls and the flood pulse on the structure of fish assemblages of the middle Xingu River in the eastern Amazon basin. Brazilian J. Biol. 75(3 suppl 1):78–94.). Ephemeroptera presents a high diversity in lotic environments, where some genera have preference for places with higher current velocity (Hamada et al. 2019HAMADA, N., NESSIMIAN, J. & QUERINO, R. 2019. Insetos Aquáticos na Amazônia brasileira: taxonomia, biologia e ecologia., Sawakuchi et al. 2015SAWAKUCHI, A.O., HARTMANN, G.A., SAWAKUCHI, H.O., PUPIM, F.N., BERTASSOLI, D.J., PARRA, M., ANTINAO, J.L., SOUSA, L.M., SABAJ PÉREZ, M.H., OLIVEIRA, P.E., SANTOS, R.A., SAVIAN, J.F., GROHMANN, C.H., MEDEIROS, V.B., MCGLUE, M.M., BICUDO, D.C. & FAUSTINO, S.B. 2015. The Volta Grande do Xingu: reconstruction of past environments and forecasting of future scenarios of a unique Amazonian fluvial landscape. Sci. Drill. 2021–32.). This ability to adapt to this type of environmental condition is related to its body structure that has as characteristic the flattened and elongated body, besides the presence of abdominal gills that aid in the displacement, most of the collected individuals are filterers and adapt easily to this type of environment, due to these characteristics occurred an expressive number of individuals obtained (Castro et al. 2013aCASTRO, D., HUGHES, R. & CALLISTO, M. 2013a. Influence of peak flow changes on the macroinvertebrate drift downstream of a Brazilian hydroelectric dam. Brazilian J. Biol. 73(4):775–782.).
During the dry season, the Xingu River provides an environment with high heterogeneity, with the presence of riffles in some localities, which will be used for fixation by organisms of the order Trichoptera (Spies et al. 2006SPIES, M.R., FROEHLICH, C.G. & KOTZIAN, C.B. 2006. Composition and diversity of Trichoptera (Insecta) larvae communities in the middle section of the Jacuí river and some tributaries, State of Rio Grande do Sul, Brazil. Iheringia. Série Zool. 96(4):389–398., Braun et al. 2014BRAUN, B.M., PIRES, M.M., KOTZIAN, C.B. & SPIES, M.R. 2014. Diversity and ecological aspects of aquatic insect communities from montane streams in southern Brazil. Acta Limnol. Bras. 26(2):186–198.). This study was carried out during the flood period characterized by high water level and uniformity in the landscape, however, even when submerged the riffles serve as shelter for the genera of the order Trichoptera, which has the capacity to build shelters with sediment and suspended material. These shelters are normally fixed to the substrate and are not easily carried by the stream, reducing the probability to found individuals of these group present in the water column (de Moor & Ivanov 2007DE MOOR, F.C. & IVANOV, V.D. 2007. Global diversity of caddisflies (Trichoptera: Insecta) in freshwater. In Freshwater Animal Diversity Assessment Springer Netherlands, Dordrecht, p.393–407.). The beginning of the dispersion movement may be related to the population density that the increase of density forcing individuals to disperse in the water column, like other organisms, aquatic or not (Munday et al. 2001MUNDAY, P.L., JONES, G.P. & CALEY, M.J. 2001. Interspecific competition and coexistence in a guild of coral-dwelling fishes. Ecology 82(8):2177–2189., Munday 2004MUNDAY, P.L. 2004. Competitive coexistence of coral-dwelling fishes: the lottery hypothesis revisited. Ecology 85(3):623–628., Yu et al. 2001YU, D.W., WILSON, H.B. & PIERCE, N.E. 2001. An empirical model of species coexistence in a spatially structured environment. Ecology 82(6):1761–1771.).
Since this relationship exists between the flow of a river and the processes of dispersion and drift, changes in the water dynamics of a river can drastically alter the distribution of aquatic insects, since these spatial patterns are directly related to the drift movement. In order to generate energy, Hydroelectric Power Plants are being built all over Brazil, however such developments cause great changes in the natural course of the river. As there is a relationship between aquatic insects and water flow, these modifications will alter the structure of communities, generating a new equilibrium in the environment (Fearnside 2016FEARNSIDE, P.M. 2016. Environmental policy in Brazilian Amazonia: Lessons from recent history. Novos Cad. NAEA 19(1):.). Understanding how the environment is in its pristine state is essential to create conservation strategies for ecosystem processes, allowing the maintenance and use of natural resources (Oldmeadow et al. 2010OLDMEADOW, D.F., LANCASTER, J. & RICE, S.P. 2010. Drift and settlement of stream insects in a complex hydraulic environment. Freshw. Biol. 55(5):1020–1035.). In addition, it is necessary to establish parameters to be used in monitoring human-modified environments (Gray et al. 2011GRAY, E.W., FUSCO, R.A., NOBLET, R. & WYATT, R.D. 2011. Comparison of Morning and Evening Larvicide Applications on Black Fly (Diptera: Simuliidae) Mortality. J. Am. Mosq. Control Assoc. 27(2):170–172., Hauer et al. 2012HAUER, C., UNFER, G., GRAF, W., LEITNER, P., ZEIRINGER, B. & HABERSACK, H. 2012. Hydro-morphologically related variance in benthic drift and its importance for numerical habitat modelling. Hydrobiologia 683(1):83–108.).
Our study demonstrates an organization of these organisms in an environment without major changes. However, the Belo Monte hydroelectric plant is being built on the Xingu River, which is modifying the original landscape of the river, in which some parts of the river were closed for the construction of a reservoir. These changes directly influence the community of aquatic insects that depend on the flow of the river. The barriers that have been created will make some communities isolated and over time this may result in changes in population genetics in these communities, as well as loss of diversity due to loss followed by non-replenishment of individuals of the species. This study will serve as a reference on how the community of aquatic insects was structured before the changes occurred in the river, being necessary the monitoring to observe how the community will be structured by the changes in the environment.
Acknowledgments
We thank the CNPq for the scholarship of Leal TB; to Eletronorte for the field support; to Federal University of Pará for the support in the production of the manuscript (06/2022 – PAPQ/PROPESP); Kostek L and Santos W for help in identifying the material; and we thank the two anonymous reviewers who evaluated the manuscript.
-
Data AvailabilitySupporting data are available at <https://doi.org/10.5281/zenodo.7186623>.
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Edited by
Associate Editor
Publication Dates
-
Publication in this collection
31 Mar 2023 -
Date of issue
2023
History
-
Received
08 Apr 2022 -
Accepted
24 Jan 2023