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Diet of Phylloicus (Trichoptera: Calamoceratidae) caddisfly larvae in forest streams of western Pará, central Brazilian Amazonia

Dieta de larvas de Phylloicus (Trichoptera: Calamoceratidae) em riachos florestais do Oeste do Pará, Amazônia central brasileira

Abstract:

Aim

The aquatic larvae of the Phylloicus (Trichoptera: Calamoceratidae) caddisflies are typical shredders. However, the trophic classification of Phylloicus has been based on the morphology and behaviour of the larvae. The aim of this study was to investigate the diet of caddisfly larvae Phylloicus in streams.

Methods

In order to provide a more reliable classification of Phylloicus diet, we analyzed the stomach contents of 185 larvae collected, sampled with D-frame entomological net from 18 streams located in the tropical forests of western Pará, Brazil. We compared the stomach contents between the larval stages, seasons (dry and rainy), and substrates (i.e., sand, leaf litter, roots, clay and mixed substrates).

Results

We identified the stomach contents as fine particulate organic matter (FPOM), coarse particulate organic matter (CPOM) and plant tissue. The diet of the Phylloicus larvae was composed basically of FPOM, independent of the larval stage (90,4%), was a higher consumption of FPOM in the dry season and there were no significant differences in food resource between substrates.

Conclusions

These findings indicate that the classification of Phylloicus as a shredder may in fact be based on the processing of leaves for the building of larval cases, rather than the diet, which is in fact detritivorous, enjoying the availability of FPOM in the streams. The great quantity of FPOM consumed by caddisfly larvae Phylloicus highlight the importance of this food resource for macroinvertebrate communities from tropical streams. Therefore, regional studies of feeding habits are needed for accurate classification trophic of Phylloicus.

Keywords:
aquatic insects; tropical streams; trophic ecology; substrates

Resumo:

Objetivo

Phylloicus (Trichoptera: Calamoceratidae) é um gênero de inseto aquático cujas larvas são consideradas fragmentadoras típicas. Porém, a classificação trófica de Phylloicus tem sido baseada na morfologia e comportamento das larvas. O objetivo deste estudo foi investigar a dieta de larvas de Phylloicus em riachos.

Métodos

Assim, para estabelecer a correta dieta alimentar de Phylloicus analisamos o conteúdo estomacal de 185 larvas coletadas, amostradas com rede entomológica D em 18 riachos de floresta tropical no Oeste do Pará, Brasil. Comparamos o conteúdo estomacal entre os estádios larvais, períodos sazonais (seco e chuvoso) e substratos (e.g. areia, folhiço, raiz, substrato misto).

Resultados

Identificamos os conteúdos estomacais como matéria orgânica particulada fina (MOPF), matéria orgânica particulada grossa (MOPG) e tecido vegetal. Independentemente do estádio larval, a dieta de Phylloicus foi composta basicamente MOPF (90,4%), houve maior consumo de MOPF no período seco e não houve diferenças significativas no consumo de recursos alimentares entre substratos.

Conclusões

Esses resultados indicam que a classificação de Phylloicus como fragmentador deve ser associada somente à quebra de folhas para a construção de abrigos e não a sua dieta, que é detritívora, aproveitando a disponibilidade de MOPF nos riachos. A grande quantidade de FPOM consumida por larvas de Phylloicus destaca a importância desse recurso alimentar para comunidades de macroinvertebrados de riachos tropicais. Portanto, estudos regionais de hábitos alimentares são necessários para uma classificação precisa trófica de Phylloicus.

Palavras-chave:
insetos aquáticos; riachos tropicais; ecologia trófica; substratos

1. Introduction

Aquatic macroinvertebrates can be classified in five basic Functional Feeding Groups (FFGs) (i.e., shredders, grazers (scrapers), collectors, filterers, and predators) based on the substrate occupied, morphological adaptations, and feeding behavior (Cummins, 2016CUMMINS, K.W. Combining taxonomy and function in the study of stream macroinvertebrates. Journal of Limnology, 2016, 75(s1), 235-241. http://dx.doi.org/10.4081/jlimnol.2016.1373.
http://dx.doi.org/10.4081/jlimnol.2016.1...
). Shredders have mouthparts adapted for the cutting and grinding of large particles of organic material, such as leaves that fall into streams from the riparian vegetation (Cushing & Allan, 2001CUSHING, C.E. and ALLAN, J.D. Streams: their ecology and life. San Diego: Academic Press, 2001.; Cobo, 2005COBO, F. Maintenance of shredders in the laboratory. In: M.A.S. GRAÇA, F. BÄRLOCHER and M.O. GESSNER, eds. Methods to study litter decomposition: a practical guide. Dordrecht: Springer, 2005, pp. 291-295. http://dx.doi.org/10.1007/1-4020-3466-0_40.
http://dx.doi.org/10.1007/1-4020-3466-0_...
). As they feed, aquatic macroinvertebrates shredders reduce the coarse particulate organic matter (CPOM) into increasingly smaller fragments (fine particulate organic matter, FPOM), enabling the transfer of nutrients and energy to other components of the trophic web (Allan & Castillo, 2007ALLAN, J.D. and CASTILLO, M.M. Stream ecology: structure and function of running Waters. Dordrecht: Springer, 2007. http://dx.doi.org/10.1007/978-1-4020-5583-6.
http://dx.doi.org/10.1007/978-1-4020-558...
; Boyero et al., 2011BOYERO, L., PEARSON, R.G., DUDGEON, D., GRAÇA, M.A., GESSNER, M.O., ALBARIÑO, R.J., FERREIRA, V., YULE, C.M., BOULTON, A.J., ARUNACHALAM, M., CALLISTO, M., CHAUVET, E., RAMÍREZ, A., CHARÁ, J., MORETTI, M.S., GONÇALVES JÚNIOR, J.F., HELSON, J.E., CHARÁ-SERNA, A.M., ENCALADA, A.C., DAVIES, J.N., LAMOTHE, S., CORNEJO, A., LI, A.O., BURIA, L.M., VILLANUEVA, V.D., ZÚÑIGA, M.C. and PRINGLE, C.M. Global distribution of a key trophic guild contrasts with common latitudinal diversity patterns. Ecology, 2011, 92(9), 1839-1848. http://dx.doi.org/10.1890/10-2244.1. PMid:21939080.
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). In turn, these insects also constitute prey for fish and other vertebrates, and are thus considered to be a key group in the ecological processes that sustain the ecosystem functioning of lower order streams (Merritt & Cummins, 1996MERRITT, R.W. and CUMMINS, K.W. An introduction to the aquatic insects of North America. Dubuque: Kendall Hunt Publishing, 1996.; Merritt et al., 2008MERRITT, R.W., CUMMINS, K.W. and BERG, M.B., eds. An introduction to the aquatic insects of America. Iowa: Kendall Hunt Publishing, 2008.).

The classification of FFGs provides an important research tool that has been used widely and refined progressively over the past 50 years (Cummins, 2016CUMMINS, K.W. Combining taxonomy and function in the study of stream macroinvertebrates. Journal of Limnology, 2016, 75(s1), 235-241. http://dx.doi.org/10.4081/jlimnol.2016.1373.
http://dx.doi.org/10.4081/jlimnol.2016.1...
), permitting the evaluation of environments through the relative proportions of the different FFGs, which also provide important insights into habitat quality (e.g., Couceiro et al., 2011COUCEIRO, S.R.M., HAMADA, N., FORSBERG, B.R. and PADOVESI-FONSECA, C. Trophic structure of macroinvertebrates in Amazonian streams impacted by anthropogenic siltation. Austral Ecology, 2011, 36, 628-637.). Despite its analytical value, the classification of FFGs should be treated with caution when applied to the assessment of habitat quality or the trophic structure of ecosystems in tropical regions, in order to avoid biased interpretations. This is because a number of studies have shown that the diet of macroinvertebrates may vary also with life instar (Malas & Wallace, 1977MALAS, D. and WALLACE, J.B. Strategies for coexistence in three species of net-spinning caddisflies (Trichoptera) in second-order southern Appalachian streams. Canadian Journal of Zoology, 1977, 55, 1829-1840.; Casas, 1996CASAS, J.J. The effect of diet quality on growth and development of recently hatched larvae of Chironomus gr. plumosus. Limnetica, 1996, 12, 1-8.; Merritt et al., 2014MERRITT, R.W., CUMMINS, K.W. and CAMPBELL, E.Y. Uma abordagem funcional para a caracterização de riachos brasileiros. In: N. HAMADA, J.L. NESSIMIAN and R.B. QUERINO, eds. Insetos aquáticos na Amazônia brasileira: taxonomia, biologia e ecologia. Manaus: INPA, 2014, pp. 69-88.), and would thus not belong to a single FFG (Tomanova et al., 2006TOMANOVA, S., GOITIA, E. and HELEŠIC, J. Trophic Levels and Functional Feeding Groups of Macroinvertebrates in Neotropical Streams. Hydrobiologia, 2006, 556(1), 251-264. http://dx.doi.org/10.1007/s10750-005-1255-5.
http://dx.doi.org/10.1007/s10750-005-125...
; Silveira-Manzotti et al., 2016SILVEIRA-MANZOTTI, B.N., MANZOTTI, A.R., CENEVIVA-BASTOS, M. and CASATTI, L. Trophic structure of macroinvertebrates in tropical pasture streams. Acta Limnologica Brasiliensia, 2016, 28(0), 1-10. http://dx.doi.org/10.1590/S2179-975X0316.
http://dx.doi.org/10.1590/S2179-975X0316...
; Tierno de Figueroa et al., 2019TIERNO DE FIGUEROA, J.M., LÓPEZ-RODRÍGUEZ, M.J. and VILLAR-ARGAIZ, M. Spatial and seasonal variability in the trophic role of aquatic insects: an assessment of functional feeding group applicability. Freshwater Biology, 2019, 64(5), 954-966. http://dx.doi.org/10.1111/fwb.13277.
http://dx.doi.org/10.1111/fwb.13277...
). In most cases, this would result in a considerable reduction in the number of typical shredders found in tropical streams (Buss et al., 2002BUSS, D.F., BAPTISTA, D.F., SILVEIRA, M.P., NESSIMIAN, J.L. and DORVILLÉ, L.F.M. Influence of water chemistry and environmental degradation on macroinvertebrate assemblages in a river basin in south-east Brazil. Hydrobiologia, 2002, 481(1/3), 125-136. http://dx.doi.org/10.1023/A:1021281508709.
http://dx.doi.org/10.1023/A:102128150870...
; Biasi et al., 2013BIASI, C., TONIN, A.M., RESTELLO, R.M. and HEPP, L.U. The colonisation of leaf litter by Chironomidae (Diptera): The influence of chemical quality and exposure duration in a subtropical stream. Limnologica, 2013, 43(6), 427-433. http://dx.doi.org/10.1016/j.limno.2013.01.006.
http://dx.doi.org/10.1016/j.limno.2013.0...
; Mesa et al., 2013MESA, L.M., REYNAGA, M.C., CORREA, M.V. and SIROMBRA, M.G. Effects of anthropogenic impacts on benthic macroinvertebrates assemblages in subtropical mountain streams. Série Zoologia, 2013, 103(4), 342-349. http://dx.doi.org/10.1590/S0073-47212013000400002.
http://dx.doi.org/10.1590/S0073-47212013...
; Gonçalves Júnior et al., 2014GONÇALVES JÚNIOR, J.F., REZENDE, R.S., GREGÓRIO, R.S. and VALENTIN, G.C. Relationship between dynamics of litterfall and riparian plant species in a tropical stream. Limnologica, 2014, 44, 40-48. http://dx.doi.org/10.1016/j.limno.2013.05.010.
http://dx.doi.org/10.1016/j.limno.2013.0...
; Graça et al., 2015GRAÇA, M.A.S., FERREIRA, W.R., FIRMIANO, K., FRANÇA, J. and CALLISTO, M. Macroinvertebrate identity, not diversity, differed across patches differing in substrate particle size and leaf litter packs in low order, tropical Atlantic forest streams. Limnetica, 2015, 34, 29-40.; Ferreira et al., 2015FERREIRA, W.R., LIGEIRO, R., MACEDO, D.R., HUGHES, R.M., KAUFMANN, P.R., OLIVEIRA, L. and CALLISTO, M. Is the diet of a typical shredder related to the physical habitat of headwater streams in the Brazilian Cerrado? Annales de Limnologie-Internatonal Journal of Limnology, 2015, 51(2), 115-122. http://dx.doi.org/10.1051/limn/2015004.
http://dx.doi.org/10.1051/limn/2015004...
; Aguiar et al., 2018AGUIAR, A.C.F., NERES-LIMA, V. and MOULTON, T.P. Relationships of shredders, leaf processing and organic matter along a canopy cover gradient in tropical streams. Journal of Limnology, 2018, 77, 109-120.). This omnivorous behavior reflects the trophic flexibility of these taxa, with the composition of the diet being adjusted to the availability of food resource (Dudgeon, 2000DUDGEON, D. The ecology of tropical Asian rivers and streams in relation to biodiversity conservation. Annual Review of Ecology and Systematics, 2000, 31(1), 239-263. http://dx.doi.org/10.1146/annurev.ecolsys.31.1.239.
http://dx.doi.org/10.1146/annurev.ecolsy...
; Tamaris-Turizo et al., 2020TAMARIS-TURIZO, C.E., PINILLA-A, G.A., GUZMÁN-SOTO, C.J. and GRANADOS-MARTÍNEZ, C.E. Assigning functional feeding groups to aquatic arthropods in a Neotropical mountain river. Aquatic Biology, 2020, 29, 45-57. http://dx.doi.org/10.3354/ab00724.
http://dx.doi.org/10.3354/ab00724...
). In other words, a given taxon may be assigned to different FFGs, depending on the sampling period (Camacho et al., 2009CAMACHO, R., BOYERO, L., CORNEJO, A., IBÁÑEZ, A. and PEARSON, R.G. Local variation in shredder distribution can explain their oversight in tropical streams. Biotropica, 2009, 41(5), 625-632. http://dx.doi.org/10.1111/j.1744-7429.2009.00519.x.
http://dx.doi.org/10.1111/j.1744-7429.20...
), because in some parts of the tropics, there is stronger seasonal variation in rainfall and subsequent runoff, which transports leaf litter and debris from riparian areas to streams (Allan & Castillo, 2007ALLAN, J.D. and CASTILLO, M.M. Stream ecology: structure and function of running Waters. Dordrecht: Springer, 2007. http://dx.doi.org/10.1007/978-1-4020-5583-6.
http://dx.doi.org/10.1007/978-1-4020-558...
).

Given this, it may be difficult to apply the FFG approach reliably in many tropical streams, where few data are available on the functional composition of the macroinvertebrate community (Boyero et al., 2009BOYERO, L., RAMIREZ, A., DUDGEON, D. and PEARSON, R.G. Are tropical streams really different? Journal of the North American Benthological Society, 2009, 28(2), 397-403. http://dx.doi.org/10.1899/08-146.1.
http://dx.doi.org/10.1899/08-146.1...
). The approach has been used successfully for some taxa, where stable isotope analysis has recently proved to be a useful tool for determining the importance of food resources for consumers, architecture and dynamics of food networks in streams (Neres-Lima et al., 2016NERES-LIMA, V., BRITO, E.F., KRSULOVIĆ, F.A.M., DETWEILER, A.M., HERSHEY, A.E. and MOULTON, T.P. High importance of autochthonous basal food source for the food web of a Brazilian tropical stream regardless of shading. International Review of Hydrobiology, 2016, 101(3-4), 132-142. http://dx.doi.org/10.1002/iroh.201601851.
http://dx.doi.org/10.1002/iroh.201601851...
; Parreira de Castro et al., 2016PARREIRA DE CASTRO, D.M., REIS DE CARVALHO, D., POMPEU, P.S., MOREIRA, M.Z., NARDOTO, G.B. and CALLISTO, M. land use influences niche size and the assimilation of resources by benthic macroinvertebrates in tropical headwater streams. PLoS One, 2016, 11(3), e0150527. http://dx.doi.org/10.1371/journal.pone.0150527. PMid:26934113.
http://dx.doi.org/10.1371/journal.pone.0...
), although not all related species (congeners in tropical and temperate areas) share the same diet (Cheshire et al., 2005CHESHIRE, K., BOYERO, L. and PEARSON, R.G. Food webs in tropical Australian streams: shredders are not scarce. Freshwater Biology, 2005, 50(5), 748-769. http://dx.doi.org/10.1111/j.1365-2427.2005.01355.x.
http://dx.doi.org/10.1111/j.1365-2427.20...
; Tomanova et al., 2006TOMANOVA, S., GOITIA, E. and HELEŠIC, J. Trophic Levels and Functional Feeding Groups of Macroinvertebrates in Neotropical Streams. Hydrobiologia, 2006, 556(1), 251-264. http://dx.doi.org/10.1007/s10750-005-1255-5.
http://dx.doi.org/10.1007/s10750-005-125...
; Chara-Serna et al., 2012; Ferreira et al., 2015FERREIRA, W.R., LIGEIRO, R., MACEDO, D.R., HUGHES, R.M., KAUFMANN, P.R., OLIVEIRA, L. and CALLISTO, M. Is the diet of a typical shredder related to the physical habitat of headwater streams in the Brazilian Cerrado? Annales de Limnologie-Internatonal Journal of Limnology, 2015, 51(2), 115-122. http://dx.doi.org/10.1051/limn/2015004.
http://dx.doi.org/10.1051/limn/2015004...
). This has led to an expansion of research on the trophic classification of macroinvertebrates in tropical areas (e.g., Cummins et al., 2005CUMMINS, K.W., MERRITT, R.W. and ANDRADE, P.C.N. The use of invertebrate functional groups to characterize ecosystem attributes in selected streams and rivers in southeast Brazil. Studies on Neotropical Fauna and Environment, 2005, 40(1), 69-90. http://dx.doi.org/10.1080/01650520400025720.
http://dx.doi.org/10.1080/01650520400025...
; Tomanova et al., 2007TOMANOVA, S., TEDESCO, P.A., CAMPERO, M., VAN DAMME, P.A., MOYA, P.A.N. and OBERDORFF, T. Longitudinal and altitudinal changes of macro-invertebrate functional feeding groups in neotropical streams: a test of the River Continuum Concept. Fundamental and Applied Limnology, 2007, 170(3), 233-241. http://dx.doi.org/10.1127/1863-9135/2007/0170-0233.
http://dx.doi.org/10.1127/1863-9135/2007...
; Príncipe et al., 2010PRÍNCIPE, R.E., GUALDONI, C.M., OBERTO, A.M., RAFFAINI, G.B. and CORIGLIANO, M.C. Spatial-temporal patterns of functional feeding groups in mountain streams of Córdoba, Argentina. Ecología Austral, 2010, 20, 257-268.; Uwadiae, 2010UWADIAE, R.E. Macroinvertebrates functional feeding groups as indices of biological assessment in a tropical aquatic ecosystem: implications for ecosystem functions. New York Science Journal, 2010, 3, 6-15.; Figueroa et al., 2011FIGUEROA, R., RODRÍGUEZ-BARRIOS, J., OSPINA-TÓRRES, R. and TURIZO-CORREA, R. Grupos funcionales alimentarios de macroinvertebrados acuáticos en el río Gaira, Colombia. Revista de Biología Tropical, 2011, 59, 1537-1552.), although a regional approach must also be developed (Ferreira et al., 2015FERREIRA, W.R., LIGEIRO, R., MACEDO, D.R., HUGHES, R.M., KAUFMANN, P.R., OLIVEIRA, L. and CALLISTO, M. Is the diet of a typical shredder related to the physical habitat of headwater streams in the Brazilian Cerrado? Annales de Limnologie-Internatonal Journal of Limnology, 2015, 51(2), 115-122. http://dx.doi.org/10.1051/limn/2015004.
http://dx.doi.org/10.1051/limn/2015004...
).

In this context, we investigated the diet of caddisfly larvae Phylloicus Müller, 1880 (Trichoptera: Calamoceratidae) in some streams of central Amazonia. This genus of aquatic insects is widely distributed in the streams of South America, and its larvae are considered to be typical leaf shredders, using leaves not only for their nutrients, but also as the raw material for the construction of their cases (Wantzen & Wagner, 2006WANTZEN, K.M. and WAGNER, R. Detritus processing by invertebrate shredders: a neotropical – temperate comparison. Journal of the North American Benthological Society, 2006, 25(1), 216-232. http://dx.doi.org/10.1899/0887-3593(2006)25[216:DPBISA]2.0.CO;2.
http://dx.doi.org/10.1899/0887-3593(2006...
). However, research suggest that Phylloicus larvae have shown plasticity in their dietary behavior (Ferreira et al., 2015FERREIRA, W.R., LIGEIRO, R., MACEDO, D.R., HUGHES, R.M., KAUFMANN, P.R., OLIVEIRA, L. and CALLISTO, M. Is the diet of a typical shredder related to the physical habitat of headwater streams in the Brazilian Cerrado? Annales de Limnologie-Internatonal Journal of Limnology, 2015, 51(2), 115-122. http://dx.doi.org/10.1051/limn/2015004.
http://dx.doi.org/10.1051/limn/2015004...
; Tamaris-Turizo et al., 2020TAMARIS-TURIZO, C.E., PINILLA-A, G.A., GUZMÁN-SOTO, C.J. and GRANADOS-MARTÍNEZ, C.E. Assigning functional feeding groups to aquatic arthropods in a Neotropical mountain river. Aquatic Biology, 2020, 29, 45-57. http://dx.doi.org/10.3354/ab00724.
http://dx.doi.org/10.3354/ab00724...
). Therefore, the present study considered the developmental instar of the larvae, the season (dry or rainy), and the relationship between the larvae and the substrate for the evaluation of dietary patterns contributing to the dataset on their trophic ecology and feeding behavior in tropical streams.

2. Material and Methods

2.1. Study area

The present study was conducted in 18 streams in western Pará (Brazil), streams are first and second order watercourses located within terra firme (unflooded) forest, distributed equally (9 in each case) in two areas– (i) the Alter do Chão Environmental Protection Area (EPA), and (ii) the Tapajós National Forest (Figure 1). The Alter do Chão EPA is located in the municipalities of Santarém and Belterra, and covers a total area of ~16 thousand hectares, limited to the north and west by the right margin of the Tapajós River, to the south by the Aramanaí EPA and Jurutui stream, and to the west by the Mojuí dos Campos land concession (Carvalho Júnior et al., 2008CARVALHO JÚNIOR, E.A.R., LIMA, A.P., MAGNUSSON, W.E. and ALBERNAZ, A.L.K.M. Long-term Effect of Forest Fragmentation on the Amazonian Gekkonid Lizards, Coleodactylus amazonicus and Gonatodes humeralis. Austral Ecology, 2008, 33(6), 723-729. http://dx.doi.org/10.1111/j.1442-9993.2008.01840.x.
http://dx.doi.org/10.1111/j.1442-9993.20...
; Vasconcelos et al., 2008VASCONCELOS, H.L., LEITE, M.F., VILHENA, J.M.S., LIMA, A.P. and MAGNUSSON, W.E. Ant diversity in an Amazonian savanna: Relationship with vegetation structure, disturbance by fire, and dominant ants. Austral Ecology, 2008, 33(2), 221-231. http://dx.doi.org/10.1111/j.1442-9993.2007.01811.x.
http://dx.doi.org/10.1111/j.1442-9993.20...
). The Tapajós National Forest (549 thousand hectares) includes parts of the municipalities of Belterra, Aveiro, Placas, and Rurópolis. This protected area is limited to the south by the Cupari River, to the west by the Tapajós River, to the east by the BR-163 federal highway and to the north, by a latitudinal parallel that links km 50 of the BR-163 highway to the Tapajós River (Santarém-Cuiabá) (Silva, 2009SILVA, M.L. A educação ambiental e suas contribuições para a sustentabilidade da região amazônica: um estudo sobre as experiências desenvolvidas na Floresta Nacional do Tapajós. Interacções, 2009, 11, 122-152.; ICMBio, 2019INSTITUTO CHICO MENDES DE CONSERVAÇÃO DA BIODIVERSIDADE – ICMBio . A floresta nacional do Tapajós [online]. Santarém: ICMBio, 2019 [viewed 2 Jan. 2019]. Available from: http://www.icmbio.gov.br/flonatapajos/
http://www.icmbio.gov.br/flonatapajos/...
). This area is dominated by dense tropical rainforest (Figure 1).

Figure 1
Location of the study streams in which the Phylloicus populations were surveyed in western Pará, Brazil.

The region’s climate is of the Ami type in the Köppen classification, with mean annual temperatures of between 25 °C and 26 °C, relative humidity of over 80% throughout the year, and mean annual precipitation of approximately 2000 mm (Silva et al., 2012SILVA, E.S., MATHIAS, C.S., LIMA, M.C.F., VEIGA JUNIOR, V.F., RODRIGUES, D.P. and CLEMENT, C.R. Análise físico-química do óleo-resina e variabilidade genética de copaíba na Floresta Nacional do Tapajós. Pesquisa Agropecuária Brasileira, 2012, 47(11), 1621-1628. http://dx.doi.org/10.1590/S0100-204X2012001100009.
http://dx.doi.org/10.1590/S0100-204X2012...
). The dry season (monthly precipitation of less than 60 mm) lasts four months.

2.2. Sampling

Data were collected during the dry season (July-September) of 2013, and the subsequent rainy season, in April and May 2014. A 50-meter transect was demarcated in each stream, with five collecting points distributed at 50-m intervals. At each point, a D-frame entomological net (30 cm wide, 500 µm mesh) was dragged three times across the bottom over a distance of one meter, from the center of the stream to the margin, to sample different substrates, such as sand, roots, submerged leaf litter, clay and mixed substrates. All samples were preserved individually, and taken to the laboratory in plastic bags filled with 96% ethanol.

2.3. Abiotic variables

Abiotic variables were measured at the first, third, and fifth sampling points in each stream transect. The current speed (m.s-1) was measured by the floater method, which consists of timing the displacement of a floating object over a known distance. Stream width and depth were measured using a surveyor’s tape or meter ruler. The main discharge (m3.s-1) was obtained from the formula: discharge = stream depth × stream width × current speed. Water temperature and dissolved oxygen were measured using a portable Oakton DO 110 oximeter, while the pH and electrical conductivity were measured buy a portable Oakton pH/con 10 m waterproof potentiometer-conductivity meter.

The opening of the canopy above each stream was estimated from photographs taken with a Canon Lens digital camera. The photographs were taken with the camera facing skywards at approximately 70 cm above the surface of the water. In the laboratory, the photographs were converted into black-and-white using the limiar function in Adobe Photoshop CS4. The percentage canopy opening was estimated from the number of white pixels in each photograph.

2.4. Processing and analysis of the stomach contents

In the laboratory, the samples were processed using a stereoscopic microscope, Petri dishes, and entomological tweezers. The Phylloicus larvae were counted for each stream and substrate, and then stored for the analysis of the stomach contents. The stomach contents were extracted from each specimen using entomological tweezers and pins, and analyzed under an optical microscope using the transparency method (Tomanova et al., 2006TOMANOVA, S., GOITIA, E. and HELEŠIC, J. Trophic Levels and Functional Feeding Groups of Macroinvertebrates in Neotropical Streams. Hydrobiologia, 2006, 556(1), 251-264. http://dx.doi.org/10.1007/s10750-005-1255-5.
http://dx.doi.org/10.1007/s10750-005-125...
). The proportion of each item in the stomach contents was estimated based on the relative area occupied by the particles at 10 points chosen randomly on each slide (Zeiss Stemi 2000, magnification of 100 x or 400 x). The proportion of each item observed at the 10 points was calculated and divided by 100. Five categories of food items — modified from Tomanova et al. (2006)TOMANOVA, S., GOITIA, E. and HELEŠIC, J. Trophic Levels and Functional Feeding Groups of Macroinvertebrates in Neotropical Streams. Hydrobiologia, 2006, 556(1), 251-264. http://dx.doi.org/10.1007/s10750-005-1255-5.
http://dx.doi.org/10.1007/s10750-005-125...
— were used (codes within parentheses): fine debris < 1 mm (FPOM), plant tissue < 1 mm (veg), coarse debris > 1 mm (CPOM), algae (alg) and invertebrates (inv).

2.5. Data analysis

2.5.1. Determination of the larval instar of Phylloicus

The width of the cephalic capsule (the distance between the two most external points) was measured to determine the larval instar (Ferreira et al., 2015FERREIRA, W.R., LIGEIRO, R., MACEDO, D.R., HUGHES, R.M., KAUFMANN, P.R., OLIVEIRA, L. and CALLISTO, M. Is the diet of a typical shredder related to the physical habitat of headwater streams in the Brazilian Cerrado? Annales de Limnologie-Internatonal Journal of Limnology, 2015, 51(2), 115-122. http://dx.doi.org/10.1051/limn/2015004.
http://dx.doi.org/10.1051/limn/2015004...
). The larval instars were defined by compiling a table of width intervals for the measurements of the cephalic capsules of the Phylloicus larvae. The validity of the instars determined from the distribution of these intervals was tested using a one-way Analysis of Variance (ANOVA), with a post hoc Tukey’s test. Overall, cephalic capsule measures and stomach contents were obtained for 185 Phylloicus larvae, including 95 during the dry season and 90 during the rainy season.

2.5.2. Variation of stomach contents between instars and between substrates

To assess difference in the composition in diet of Phylloicus between the larval instars (proportion of each item in each size instar) and difference in diet between substrates, we conducted a one-way ANOVA, followed by a post-hoc Tukey’s test.

2.5.3. Variation of abiotic variables, food resource and Phylloicus larvae

To determine whether the abiotic variables, as well as the quantities of FPOM, CPOM and leaf matter in the stomach contents was significantly different between seasons, were applied a paired t-test.

Pearson correlation coefficients were used to evaluate the relationships between the abiotic variables and the abundance of Phylloicus larvae.

All the analyses were run in STATISTICA (v. 10.0).

3. Results

3.1. Abiotic variables

The study streams had low pH (4.83) and electrical conductivity (17.42 µS.cm-1) and moderate concentrations of dissolved oxygen (5 mg. L-1). Only three (stream depth, width, and current speed) of the eight abiotic variables analyzed varied significantly between seasons, and all returned higher values during the rainy season. (Table 1).

Table 1
Comparison of the abiotic variables recorded between the dry (07-09/2013) and rainy (04-05/2014) seasons.

3.2. Phylloicus and stomach contents

The cephalic capsules of the Phylloicus larvae varied in width from 0.25 to 2.63 mm (mean 0.96 mm ± 0.42). The larvae were distributed in five larval instars (ANOVA F(4, 180) = 379.85, p < 0.01), with 53 larvae in instar I (0.25-0.73 mm), 86 in instar II (0.74-1.22 mm), 39 in instar III (1.23-1.71 mm), five in instar IV (1.72-2.20 mm), and two in instar V (≥ 2.21 mm).

The FPOM was the predominant item in the stomach contents of the Phylloicus larvae irrespective of the larval instar, but no significant variation was found (F(4, 180) = 1.28, p = 0.28). Even if only the fraction of plant tissue is included in the analysis, no significant variation was found among the different larval instars (F(4, 180) = 1.15, p = 0.33).The CPOM was found in the stomach contents of only four specimens (2% of the larvae). However, the quantity of CPOM varied significantly among the different larval instars (F(4, 180) = 6.37, p < 0.01; Figure 2), being significantly more abundant in the final instar, albeit with reduced abundance. Based on these findings, Phylloicus was classified as a detritivore, even though it is classified as a typical shredder in the current literature.

Figure 2
Variation in the proportion of fine particulate organic matter (FPOM) (a), plant tissue (b) and coarse particulate organic matter (CPOM) (c) ingested by the different larval instars of Phylloicus collected from forest streams in western Pará, Brazil.

The abundance of Phylloicus was related significantly to the sites with low current speeds and lower concentrations of dissolved oxygen (Table 2), as found in the calm backwaters of the streams, where larger quantities of leaf litter tend to accumulate. Despite the greater abundance of Phylloicus in this type of environment, the ingestion of FPOM varied little among substrates (F(4, 180) = 1.98, p = 0.09; Figure 3). The ingestion of plant tissue was also similar among the different substrates, including those with no leaf litter (F(4, 180) = 2.01, p = 0.09). There was also little variation among substrates in the consumption of CPOM (F(4, 180) = 1.58, p = 0.18). Significantly larger quantities of FPOM were ingested during the dry season in comparison with the rainy season (t = 4.95, p= < 0.01, df = 183), although no significant seasonal variation was observed in the ingestion of CPOM (t = 0.95, p = 0.34, df = 183). However, the ingestion of plant tissue, which is a component of the CPOM, was significantly higher during the rainy season (t = -5.11, p < 0.01, df = 183).

Table 2
Pearson correlations between the abiotic variables and the abundance of Phylloicus larvae in the streams in western Pará, Brazil.
Figure 3
Variation among substrates in the proportion of fine particulate organic matter (FPOM) (a), plant tissue (b) and coarse particulate organic matter (CPOM) (c) found in the stomach contents of the Phylloicus larvae collected from forest streams in western Pará, Brazil.

4. Discussion

The classification of Phylloicus as a shredder is based on the morphology of the mouthparts and the behavior of the larvae of this genus. Phylloicus larvae cut pieces of leaves with their masticatory apparatus to build their cases (Merritt & Cummins, 1996MERRITT, R.W. and CUMMINS, K.W. An introduction to the aquatic insects of North America. Dubuque: Kendall Hunt Publishing, 1996.; Merritt et al., 2008MERRITT, R.W., CUMMINS, K.W. and BERG, M.B., eds. An introduction to the aquatic insects of America. Iowa: Kendall Hunt Publishing, 2008.). The association of these larvae with banks of leaf litter is probably advantageous in terms of protection from predators by providing camouflage and cover, but does not necessarily imply the consumption of CPOM. In this case, the shredding behavior of these larvae would represent a behavioral (predator avoidance) strategy, rather than a feeding adaptation (Cerezer et al., 2016CEREZER, C., BIASI, C., COGO, G.B. and SANTOS, S. Avoid predation or take risks in basic activities? Predator–prey relationship in subtropical streams between decapods and caddisflies. Marine and Freshwater Research, 2016, 67(12), 1880. http://dx.doi.org/10.1071/MF15278.
http://dx.doi.org/10.1071/MF15278...
).

A number of studies have shown that the composition of the diet of aquatic invertebrates varies among the different larval instars (Casas, 1996CASAS, J.J. The effect of diet quality on growth and development of recently hatched larvae of Chironomus gr. plumosus. Limnetica, 1996, 12, 1-8.; Merritt et al., 2014MERRITT, R.W., CUMMINS, K.W. and CAMPBELL, E.Y. Uma abordagem funcional para a caracterização de riachos brasileiros. In: N. HAMADA, J.L. NESSIMIAN and R.B. QUERINO, eds. Insetos aquáticos na Amazônia brasileira: taxonomia, biologia e ecologia. Manaus: INPA, 2014, pp. 69-88.). In the present study, FPOM was the predominant item in the stomach contents of the Phylloicus specimens, irrespective of the larval instar, which is consistent with the findings of Ferreira et al. (2015)FERREIRA, W.R., LIGEIRO, R., MACEDO, D.R., HUGHES, R.M., KAUFMANN, P.R., OLIVEIRA, L. and CALLISTO, M. Is the diet of a typical shredder related to the physical habitat of headwater streams in the Brazilian Cerrado? Annales de Limnologie-Internatonal Journal of Limnology, 2015, 51(2), 115-122. http://dx.doi.org/10.1051/limn/2015004.
http://dx.doi.org/10.1051/limn/2015004...
. Ecosystems such as streams receive the input of large quantities of particulate organic matter from the erosion of the margins. The vegetation that falls into tropical streams is decomposed into fine detritus very rapidly (Mathuriau & Chauvet, 2002MATHURIAU, C. and CHAUVET, E. Breakdown of leaf litter in a neotropical stream. Journal of the North American Benthological Society, 2002, 21(3), 384-396. http://dx.doi.org/10.2307/1468477.
http://dx.doi.org/10.2307/1468477...
; Gonçalves Júnior et al., 2012GONÇALVES JÚNIOR, J.F., REZENDE, R.S., MARTINS, R.S.N.M. and GREGÓRIO, R.S. Leaf breakdown in an Atlantic Rain Forest. Austral Ecology, 2012, 37(7), 807-815. http://dx.doi.org/10.1111/j.1442-9993.2011.02341.x.
http://dx.doi.org/10.1111/j.1442-9993.20...
), and continuously throughout the year. In this case, the presence of FPOM (detritus) in the stomach contents of the Phylloicus larvae may be related primarily to the availability of this material in the habitat (Henriques-Oliveira et al., 2003HENRIQUES-OLIVEIRA, A.L., NESSIMIAN, J.L. and DORVILLÉ, J.F.M. Feeding habits of Chironomid larvae (Insecta: Diptera) from a stream in the Floresta da Tijuca, Rio de Janeiro, Brazil. Brazilian Journal of Biology = Revista Brasileira de Biologia, 2003, 63(2), 269-281. http://dx.doi.org/10.1590/S1519-69842003000200012. PMid:14509849.
http://dx.doi.org/10.1590/S1519-69842003...
; Chará-Serna et al., 2012CHARÁ-SERNA, A.M., CHARA, J.D., ZUNIGA, M.D., PEARSON, R.G. and BOYERO, L. Diets of leaf litter-associated invertebrates in three tropical streams. International Journal of Limnology, 2012, 48(2), 139-144. http://dx.doi.org/10.1051/limn/2012013.
http://dx.doi.org/10.1051/limn/2012013...
). This reinforces the importance of this food resource in tropical freshwater ecosystems (Palmer et al., 1993PALMER, C., O’KEEFFE, J., PALMER, A., DUNNE, T. and RADLOFF, S. Macroinvertebrate functional feeding groups in the middle and lower reaches of the Buffalo River eastern Cape, South Africa. I. Dietary variability. Freshwater Biology, 1993, 29(3), 441-453. http://dx.doi.org/10.1111/j.1365-2427.1993.tb00778.x.
http://dx.doi.org/10.1111/j.1365-2427.19...
; Motta & Uieda., 2004MOTTA, R.L. and UIEDA, V.S. Diet and trophic groups of an aquatic insect community in a tropical stream. Brazilian Journal of Biology = Revista Brasileira de Biologia, 2004, 64(4), 809-817. http://dx.doi.org/10.1590/S1519-69842004000500010. PMid:15744421.
http://dx.doi.org/10.1590/S1519-69842004...
; Ceneviva-Bastos & Casatti, 2014CENEVIVA-BASTOS, M. and CASATTI, L. Shading effects on community composition and food web structure of a deforested pasture stream: evidences from a field experiment in Brazil. Limnologica, 2014, 46, 9-21. http://dx.doi.org/10.1016/j.limno.2013.11.005.
http://dx.doi.org/10.1016/j.limno.2013.1...
; Silveira-Manzotti et al., 2016SILVEIRA-MANZOTTI, B.N., MANZOTTI, A.R., CENEVIVA-BASTOS, M. and CASATTI, L. Trophic structure of macroinvertebrates in tropical pasture streams. Acta Limnologica Brasiliensia, 2016, 28(0), 1-10. http://dx.doi.org/10.1590/S2179-975X0316.
http://dx.doi.org/10.1590/S2179-975X0316...
), related to microbial activity, which accelerates decomposition, increasing the nutritional quality of this detritus for these invertebrates (Robinson et al., 1998ROBINSON, C.T., GESSNER, M.O. and WARD, J.V. Leaf breakdown and associated macroinvertebrates in alpine glacial streams. Freshwater Biology, 1998, 40(2), 215-228. http://dx.doi.org/10.1046/j.1365-2427.1998.00343.x.
http://dx.doi.org/10.1046/j.1365-2427.19...
; Graça, 2001GRAÇA, M.A.S. The Role of Invertebrates on Leaf Litter Decomposition in Streams – a Review. International Review of Hydrobiology, 2001, 86(4-5), 383-393. http://dx.doi.org/10.1002/1522-2632(200107)86:4/5<383::AID-IROH383>3.0.CO;2-D.
http://dx.doi.org/10.1002/1522-2632(2001...
).

Abiotic factors, such as the type of substrate or current speed influence the distribution of aquatic insects (Fidelis et al., 2008FIDELIS, L., NESSIMIAN, J.L. and HAMADA, N. Distribuição espacial de insetos aquáticos em igarapés de pequena ordem na Amazônia Central. Acta Amazonica, 2008, 38(1), 127-134. http://dx.doi.org/10.1590/S0044-59672008000100014.
http://dx.doi.org/10.1590/S0044-59672008...
) and the exploitation of food resource (Ferreira et al., 2015FERREIRA, W.R., LIGEIRO, R., MACEDO, D.R., HUGHES, R.M., KAUFMANN, P.R., OLIVEIRA, L. and CALLISTO, M. Is the diet of a typical shredder related to the physical habitat of headwater streams in the Brazilian Cerrado? Annales de Limnologie-Internatonal Journal of Limnology, 2015, 51(2), 115-122. http://dx.doi.org/10.1051/limn/2015004.
http://dx.doi.org/10.1051/limn/2015004...
). These abiotic characteristics may thus be considered predictors of the foraging behavior of aquatic macroinvertebrates. Tomanova et al. (2006)TOMANOVA, S., GOITIA, E. and HELEŠIC, J. Trophic Levels and Functional Feeding Groups of Macroinvertebrates in Neotropical Streams. Hydrobiologia, 2006, 556(1), 251-264. http://dx.doi.org/10.1007/s10750-005-1255-5.
http://dx.doi.org/10.1007/s10750-005-125...
, for example, suggested that the feeding behavior of typical shredders may shift to detritivore (collector behavior) as current speeds increase, leading to a growth in the decomposition of the leaf litter, caused by the rise in the physical abrasion of the leaves, leading to their mechanical fragmentation (Abelho, 2001ABELHO, M. From litterfall to breakdown in streams: a review. The Scientific World Journal, 2001, 1, 656-680. http://dx.doi.org/10.1100/tsw.2001.103. PMid:12805769.
http://dx.doi.org/10.1100/tsw.2001.103...
), which probably imposes energetic restrictions for the harvesting of CPOM, and an increase in the availability of fine detritus.

In the present study, the abundance of Phylloicus was related significantly to areas with the lower current speeds and lower dissolved oxygen concentrations that characterize backwater environments. This provides indirect evidence that the preference for FPOM is not influenced by any increase in current speed, but rather to other factors, such as the reduced nutritional quality of the leaves (Boyero et al., 2011BOYERO, L., PEARSON, R.G., DUDGEON, D., GRAÇA, M.A., GESSNER, M.O., ALBARIÑO, R.J., FERREIRA, V., YULE, C.M., BOULTON, A.J., ARUNACHALAM, M., CALLISTO, M., CHAUVET, E., RAMÍREZ, A., CHARÁ, J., MORETTI, M.S., GONÇALVES JÚNIOR, J.F., HELSON, J.E., CHARÁ-SERNA, A.M., ENCALADA, A.C., DAVIES, J.N., LAMOTHE, S., CORNEJO, A., LI, A.O., BURIA, L.M., VILLANUEVA, V.D., ZÚÑIGA, M.C. and PRINGLE, C.M. Global distribution of a key trophic guild contrasts with common latitudinal diversity patterns. Ecology, 2011, 92(9), 1839-1848. http://dx.doi.org/10.1890/10-2244.1. PMid:21939080.
http://dx.doi.org/10.1890/10-2244.1...
) and their rigidity, which may influence the exploitation of CPOM more than the abiotic variables measured in the study.

The type of substrate has also been used as a predictor of the abundance and diversity of macroinvertebrates (Minshall, 1984MINSHALL, G.W. Aquatic insect-substratum relationships. In: D.M. RESH and V.H. ROSEMBERG, eds. The ecology of aquatic insects. New York: Praeger Scientific, 1984, pp. 358-400.). Fidelis et al. (2008)FIDELIS, L., NESSIMIAN, J.L. and HAMADA, N. Distribuição espacial de insetos aquáticos em igarapés de pequena ordem na Amazônia Central. Acta Amazonica, 2008, 38(1), 127-134. http://dx.doi.org/10.1590/S0044-59672008000100014.
http://dx.doi.org/10.1590/S0044-59672008...
confirmed that many taxa were more abundant in given types of substrate than others, a pattern related to the characteristics of the substrate and the type of feeding resource it contains. In the present study, however, the Phylloicus larvae ingested similar amounts of FPOM in practically all the substrates. In this case, the preference of the larvae for a given substrate may be more closely related to the availability of materials for building their cases, given that the large banks of leaf litter found in first order streams are appropriate for the development of Phylloicus larvae (Landeiro et al., 2010LANDEIRO, V.L., HAMADA, N., GODOY, B.S. and MELO, A.S. Effects of litter patch area on macroinvertebrate assemblage structure and leaf breakdown in Central Amazonian streams. Hydrobiologia, 2010, 649(1), 355-363. http://dx.doi.org/10.1007/s10750-010-0278-8.
http://dx.doi.org/10.1007/s10750-010-027...
).

The FPOM was ingested in significantly larger quantities during the dry season in comparison with the rainy season, although the ingestion of CPOM did not increase during the rainy season. This may reflect the difficulties faced by the larvae for the acquisition of nutrients during a period when the environment is less stable, with the rains lead to an increase in stream depth and width, and current speeds, all of which result in the removal of food resource (Silva et al., 2009SILVA, F.L., PAULETO, G.M., TALAMONI, J.L.B. and RUIZ, S.S. Categorização funcional trófica das comunidades de macroinvertebrados de dois reservatórios na região Centro-Oeste do Estado de São Paulo, Brasil. Acta Scientiarum. Biological Sciences, 2009, 31(1), 73-78. http://dx.doi.org/10.4025/actascibiolsci.v31i1.331.
http://dx.doi.org/10.4025/actascibiolsci...
). While the ingestion of CPOM was low in both seasons, the ingestion of leaf tissue increased during the rainy season, which may be related to an increase in the input of allochthonous resources, such as fragments of plants, which are transferred to the streams by the rain (Allan & Castillo, 2007ALLAN, J.D. and CASTILLO, M.M. Stream ecology: structure and function of running Waters. Dordrecht: Springer, 2007. http://dx.doi.org/10.1007/978-1-4020-5583-6.
http://dx.doi.org/10.1007/978-1-4020-558...
; Uieda & Motta, 2007UIEDA, V.S. and MOTTA, R.L. Trophic organization and food web structure of southeastern Brazilian streams: a review. Acta Limnologica Brasiliensia, 2007, 19, 15-30.; Carvalho & Uieda, 2009CARVALHO, E.M. and UIEDA, V.S. Diet of invertebrates sampled in leaf-bags incubated in a tropical headwater stream. Zoologia, 2009, 26(4), 694-704. http://dx.doi.org/10.1590/S1984-46702009000400014.
http://dx.doi.org/10.1590/S1984-46702009...
; Lisboa et al., 2015LISBOA, L.K., SILVA, A.L.L., SIEGLOCH, A.E., GONÇALVES JÚNIOR, J.F. and PETRUCIO, M.M. Temporal dynamics of allochthonous coarse particulate organic matter in a subtropical Atlantic rainforest Brazilian stream. Marine and Freshwater Research, 2015, 66(8), 674-680. http://dx.doi.org/10.1071/MF14068.
http://dx.doi.org/10.1071/MF14068...
). This increase in plant material implies that a greater diversity of plant species becomes available, increasing the probability that palatable items will be encountered (Lima & Gonçalves, 2015LIMA, L.S. and GONÇALVES, J.F.J. Heterogeneidade temporal e espacial na composição química do detrito foliar. In: Livro de Resumos do II Simpósio Processos Ecológicos, Restauração e Ecovaloração em Zonas Ripárias (AquaRipária). Brasília: AquaRipária, 2015, pp. 12-15.), with their exploitation by the larvae accelerating the decomposition of the material into fine detritus, as mentioned above.

Analyzes of stomach contents and stable isotopes in a number of tropical streams have shown that, despite the presence of dense riparian forest and the large amounts of organic material derived from the forest, the resident invertebrates satisfy most of their energetic requirements through the exploitation of autochthonous resources (Brito et al., 2006BRITO, E.F., MOULTON, T.P., SOUZA, M.L. and BUNN, S. Stable isotope analysis indicates microalgae as the predominant food source of fauna in a coastal forest stream, southeast Brazil. Austral Ecology, 2006, 31(5), 623-633. http://dx.doi.org/10.1111/j.1442-9993.2006.01610.x.
http://dx.doi.org/10.1111/j.1442-9993.20...
; Lau et al., 2008LAU, D.C.P., LEUNG, K.M.Y. and DUDGEON, D. Experimental dietary manipulations for determining the relative importance of allochthonous and autochthonous food resources in tropical streams. Freshwater Biology, 2008, 53, 139-147.; Dudgeon et al., 2010DUDGEON, D., CHEUNG, F.K.W. and MANTEL, S.K. Food web structure in small streams: do we need different models for the tropics? Journal of the North American Benthological Society, 2010, 29(2), 395-412. http://dx.doi.org/10.1899/09-058.1.
http://dx.doi.org/10.1899/09-058.1...
; Neres-Lima et al., 2016NERES-LIMA, V., BRITO, E.F., KRSULOVIĆ, F.A.M., DETWEILER, A.M., HERSHEY, A.E. and MOULTON, T.P. High importance of autochthonous basal food source for the food web of a Brazilian tropical stream regardless of shading. International Review of Hydrobiology, 2016, 101(3-4), 132-142. http://dx.doi.org/10.1002/iroh.201601851.
http://dx.doi.org/10.1002/iroh.201601851...
). Given this, some authors have suggested that Neotropical shredders are more generalist than the species found in temperate zones (Wantzen & Wagner, 2006WANTZEN, K.M. and WAGNER, R. Detritus processing by invertebrate shredders: a neotropical – temperate comparison. Journal of the North American Benthological Society, 2006, 25(1), 216-232. http://dx.doi.org/10.1899/0887-3593(2006)25[216:DPBISA]2.0.CO;2.
http://dx.doi.org/10.1899/0887-3593(2006...
). It is also important to remember that many aquatic insects are not restricted to a single functional feeding group, as in the case of the chironomid larvae (Nessimian & Carvalho, 1998NESSIMIAN, J.L. and CARVALHO, A.L. Ecologia de insetos aquáticos. Rio de Janeiro: Programa de Pós-graduação em Ecologia, Universidade Federal do Rio de Janeiro, 1998.; Nessimian et al., 1999NESSIMIAN, J.L., SANVERINO, A.M. and OLIVEIRA, A. Relações tróficas de larvas de Chironomidae (Diptera) e sua importância na rede alimentar em um brejo de dunas no Estado do Rio de Janeiro. Revista Brasileira de Entomologia, 1999, 43, 47-53.; Henriques-Oliveira et al., 2003HENRIQUES-OLIVEIRA, A.L., NESSIMIAN, J.L. and DORVILLÉ, J.F.M. Feeding habits of Chironomid larvae (Insecta: Diptera) from a stream in the Floresta da Tijuca, Rio de Janeiro, Brazil. Brazilian Journal of Biology = Revista Brasileira de Biologia, 2003, 63(2), 269-281. http://dx.doi.org/10.1590/S1519-69842003000200012. PMid:14509849.
http://dx.doi.org/10.1590/S1519-69842003...
) or shredders in general (Mihuc & Mihuc, 1995MIHUC, T.B. and MIHUC, J.R. Trophic ecology of five shredders in a Rocky Mountain stream. Journal of Freshwater Ecology, 1995, 10(3), 209-216. http://dx.doi.org/10.1080/02705060.1995.9663440.
http://dx.doi.org/10.1080/02705060.1995....
; Rosi-Marshall & Wallace, 2002ROSI-MARSHALL, E.J. and WALLACE, B. Invertebrate food webs along a stream resource gradient. Freshwater Biology, 2002, 47(1), 129-141. http://dx.doi.org/10.1046/j.1365-2427.2002.00786.x.
http://dx.doi.org/10.1046/j.1365-2427.20...
; Ferreira et al., 2015FERREIRA, W.R., LIGEIRO, R., MACEDO, D.R., HUGHES, R.M., KAUFMANN, P.R., OLIVEIRA, L. and CALLISTO, M. Is the diet of a typical shredder related to the physical habitat of headwater streams in the Brazilian Cerrado? Annales de Limnologie-Internatonal Journal of Limnology, 2015, 51(2), 115-122. http://dx.doi.org/10.1051/limn/2015004.
http://dx.doi.org/10.1051/limn/2015004...
). These studies have emphasized the importance of stomach contents analyzes for the reliable classification of functional feeding groups, which are classified based solely on their morphological and behavioral traits.

The results of the present study reinforce the importance of data on the stomach contents of aquatic macroinvertebrates, not only in the Amazon basin, but in all tropical aquatic ecosystems, for the reliable characterization of the role of the taxa in the trophic ecology of streams. These data will provide a much more systematic understanding of the structure and trophic dynamics of these aquatic ecosystems.

Acknowledgements

The authors are grateful to the Brazilian National Council for Science and Technology (CNPq) for providing financial support for the projects “Trophic categorization of aquatic insects using morphological traits, stomach contents, and stable isotopes” (Universal/CNPq process 477187/2012-9) and “Functional trophic groups of aquatic insects in Amazonian streams in different habitats, with emphasis on the shredders” (Bionorte/CNPq process 407698/2013-2).

  • Cite as: Pimentel, D.R., Couceiro, S.R.M. and Salcedo, A.K.M. Diet of Phylloicus (Trichoptera: Calamoceratidae) caddisfly larvae in forest streams of western Pará, central Brazilian Amazonia. Acta Limnologica Brasiliensia, 2020, vol. 32, e13.

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Edited by

Associate Editors: André Megali Amado.

Publication Dates

  • Publication in this collection
    01 July 2020
  • Date of issue
    2020

History

  • Received
    03 Jan 2019
  • Accepted
    11 May 2020
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