Trophic functioning of macrobenthic fauna in a tropical acidified Bornean estuary(Southeast Asia)
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摘要
The trophic structure of a community is used to infer ecosystem functioning(e.g. energy transfer and nutrient cycling). Here the trophic structure of the benthic infaunal and epifaunal communities in the Brunei Estuary are characterized, and their distribution along an estuarine pH gradient is analyzed using univariate and multivariate techniques. This analysis revealed that surface deposit feeders(e.g., polychaetes) were numerically dominant within the infaunal communities whereas in the epifaunal communities filter feeders(e.g., bivalves) were highly abundant. Species richness for almost all trophic groups increased toward the lower estuary, except for omnivores in the epifaunal communities, which decreased markedly. Both Analysis of Variance(ANOVA) and Analysis of Similarities(ANOSIM) detected significant differences in the density of respective trophic groups among stations. Within infaunal communities, both Biological and Environmental procedure(BIO-ENV) and Canonical Correspondence Analysis(CCA) showed that trophic shifts were associated with environmental gradients. Surface-deposit feeders and omnivores were the most abundant macrobenthos of the upper estuary characterized by low salinity, low pH, and a higher percentage of mud particles. The proportion of filter feeders and carnivores increased with salinity/pH and sand. A more uniform distribution of trophic structure was found in the lower estuary, with high salinity and pH over sandy habitat. In contrast, within epifaunal trophic groups,the percentage of surface deposit feeders and omnivores declined, but filter feeders remarkably increased toward the sea. The proportion of carnivores remained similar at all stations. Non-Metric Multidimensional Scaling(nMDS) ordination for epifaunal trophic groups clearly demarcated higher salinity/pH stations from lower salinity/pH stations, suggesting different trophic compositions along the estuarine pH gradient.
The trophic structure of a community is used to infer ecosystem functioning(e.g. energy transfer and nutrient cycling). Here the trophic structure of the benthic infaunal and epifaunal communities in the Brunei Estuary are characterized, and their distribution along an estuarine pH gradient is analyzed using univariate and multivariate techniques. This analysis revealed that surface deposit feeders(e.g., polychaetes) were numerically dominant within the infaunal communities whereas in the epifaunal communities filter feeders(e.g., bivalves) were highly abundant. Species richness for almost all trophic groups increased toward the lower estuary, except for omnivores in the epifaunal communities, which decreased markedly. Both Analysis of Variance(ANOVA) and Analysis of Similarities(ANOSIM) detected significant differences in the density of respective trophic groups among stations. Within infaunal communities, both Biological and Environmental procedure(BIO-ENV) and Canonical Correspondence Analysis(CCA) showed that trophic shifts were associated with environmental gradients. Surface-deposit feeders and omnivores were the most abundant macrobenthos of the upper estuary characterized by low salinity, low pH, and a higher percentage of mud particles. The proportion of filter feeders and carnivores increased with salinity/pH and sand. A more uniform distribution of trophic structure was found in the lower estuary, with high salinity and pH over sandy habitat. In contrast, within epifaunal trophic groups,the percentage of surface deposit feeders and omnivores declined, but filter feeders remarkably increased toward the sea. The proportion of carnivores remained similar at all stations. Non-Metric Multidimensional Scaling(nMDS) ordination for epifaunal trophic groups clearly demarcated higher salinity/pH stations from lower salinity/pH stations, suggesting different trophic compositions along the estuarine pH gradient.
The trophic structure of a community is used to infer ecosystem functioning(e.g. energy transfer and nutrient cycling). Here the trophic structure of the benthic infaunal and epifaunal communities in the Brunei Estuary are characterized, and their distribution along an estuarine pH gradient is analyzed using univariate and multivariate techniques. This analysis revealed that surface deposit feeders(e.g., polychaetes) were numerically dominant within the infaunal communities whereas in the epifaunal communities filter feeders(e.g., bivalves) were highly abundant. Species richness for almost all trophic groups increased toward the lower estuary, except for omnivores in the epifaunal communities, which decreased markedly. Both Analysis of Variance(ANOVA) and Analysis of Similarities(ANOSIM) detected significant differences in the density of respective trophic groups among stations. Within infaunal communities, both Biological and Environmental procedure(BIO-ENV) and Canonical Correspondence Analysis(CCA) showed that trophic shifts were associated with environmental gradients. Surface-deposit feeders and omnivores were the most abundant macrobenthos of the upper estuary characterized by low salinity, low pH, and a higher percentage of mud particles. The proportion of filter feeders and carnivores increased with salinity/pH and sand. A more uniform distribution of trophic structure was found in the lower estuary, with high salinity and pH over sandy habitat. In contrast, within epifaunal trophic groups,the percentage of surface deposit feeders and omnivores declined, but filter feeders remarkably increased toward the sea. The proportion of carnivores remained similar at all stations. Non-Metric Multidimensional Scaling(nMDS) ordination for epifaunal trophic groups clearly demarcated higher salinity/pH stations from lower salinity/pH stations, suggesting different trophic compositions along the estuarine pH gradient.
引文
Amaral, V.. Cabral, H. N.,&Bishop. M. J.(2011). Resistance among wild invertebrate populations to recurrent estuarine acidification. Estuarine. Coastal and Shelf Science, 93(4), 460-467.
Amaral, V., Cabral, H. N.,&Bishop, M. J.(2014). Prior exposure influences the behavioural avoidance by an intertidal gastropod, Bembicium auratum, of acidified waters. Estuarine, Coastal and Shelf Science, 136, 82-90.
Bessa, F., Cunha, D., Concalves, S. C.,&Marques, J. C.(2013). Sandy beach macrofaunal assemblages as indicators of anthropogenic impacts on coastal dunes.Ecological Indicators, 30, 196-204.
Braeckman, U., Provoost, P., Gribsholt, B., van Gansbeke, D., Middelburg, J. J., Soetaert, K.,&Vanaverbeke, J.(2009). Role of macrofauna functional traits and density in biogeochemical fluxes and bioturbation. Marine Ecology Progress Series, 399, 173-186.
Brown, S. S., Gaston, G. R., Rakocinski, C. F..&Heard, R. W.(2000). Effects of sediment contaminants and environmental gradients on macrobenthic community trophic structure in Gulf of Mexico estuaries. Estuaries, 23(3), 411-424.
Carrasco, F. D.,&Carbajal, W.(1998). The distribution of polychaete feeding guilds in organic enriched sediments of San Vincente Bay, Central Chile. International Review of Hydrobiology,83,233-249.
Carstensen, J., Conley, D. J., Bonsdorff, E., Gustafsson, B. G., Hietanen, S., Janas, U.,&Voss, M.(2014). Hypoxia in the Baltic Sea:Biogeochemical cycles, benthic fauna, and management. Ambio, 43(1), 26-36.
Chardy, P.,&Clavier, J.(1988). Biomass and trophic structure of the macrobenthos in the south-west lagoon of New Caledonia. Marine Biology, 99, 105-202.
Clarke, K. R.(1993). Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology, 18, 117-143.
Clarke, K. R.,&Warwick, R. M.(2001). Change in marine communities:An approach to statistical analysis and interpretation. Plymouth, UK:Plymouth Marine Laboratory.
Dauvin, J. C.,&Desroy, N.(2005). The food web in the lower part of the Seine estuary:A synthesis of existing knowledge. Hydrobiologia, 540(1-3), 13-27.
Dauwe, B., Herman, P. M. J.,&Heip, C. H. R.(1998). Community structure and bioturbation potential of macrofauna at four North Sea stations with contrasting food supply. Marine Ecology Progress Series, 173, 67-83.
Diaz, R. J.,&Schaffner, L C.(1990). The functional role of estuarine benthos In:M. Haired,&E. C. Krome(Eds.), Perspectives on the Chesapeake Bay, 1990Advances in Estuarine Sciences(pp. 25-56). U.S. Environmental Protection Agency, Chesapeake Bay Program(Report CBP/TRS/41/90).
Dimitriadis, C.,&Koutsoubas, D.(2011). Functional diversity and species turnover of benthic invertebrates along a local environmental gradient induced by an aquaculture unit:The contribution of species dispersal ability and rarity.Hydrobiologia, 670(1), 307-315.
Dove, C. M.,&Sammut,J.(2007). Impacts of estuarine acidification on survival and growth of Sydney rock oysters Saccostrea glomerata(Gould 1850).Journal of Shellfish Research, 26, 519-527.
Duan, X. H., Wang, Z. Y., Xu, M. Z.,&Zhang, K.(2009). Effect of streambed sediment on benthic ecology. International Journal of Sediment Research, 24(3), 325-338.
Fauchald, K.,&Jumars, P.(1979). The diet of worms:A study of polychaete feeding guilds. Oceanography and Marine Biology:An annual Review, 17,194-284.
Garcia-Arberas, L,&Rallo, A.(2002). The intertidal soft bottom infaunal macrobenthos in three Basque estuaries(Gulf of Biscay):A feeding guild approach.Hydrobiologia, 475-476, 457-468.
Gaston, G. R., Brown, S. S., Rakocinski, C. F., Heard, R. W.,&Summers, J. K.(1995).Trophic structure of macrobenthic communities in Northern Gulf of Mexico estuaries. Gulf Research Reports, 9,111-116.
Gaston, G. R., Rakocinski, C. F., Brown, S. S.,&Cleveland, C. M.(1998). Trophic function in estuaries:Response of macrobenthos to natural and contaminant gradients. Marine and Freshwater Research, 49(8), 833-846.
Gaudencio, M. J.,&Cabral, H. N.(2007). Trophic structure of macrobenthos in the Tagus estuary and adjacent coastal shelf. Hydrobiologia, 587(1), 241-251.
Gray, J. S.,&Elliott, M.(2009). Ecology of marine sediments:Science to management(2nd edition). Oxford, UK:Oxford University Press.
Grilo, T. F., Cardoso, P. G., Dolbeth, M., Bordalo, M. D.,&Pardal, M. A.(2011). Effects of extreme climate events on the macrobenthic communities'structure and functioning of a temperate estuary. Marine Pollution Bulletin, 62(2), 303-311.
Hammer, 0., Harper, D. A.,&Ryan, P. D.(2001). PAST:Paleontological statistics software package for education and data analysis. Palaeontologia Electronica, 4(1), 9.
Hanekom, N.(2011). Trophic structure and biomass distribution of macrobenthos on sheltered and semi-exposed rocky shores of Tsitsikamma Marine Protected Area. African Zoology, 46(2), 224-238.
Hooi, S. M. D.(1987). Sungai Brunei water quality and pollution study(Final Report).Brunei Darussalam:Government of Negara Brunei Darussalam.
Hooper, D. U., Chapin, F. S.,Ⅲ, Ewel, J. J., Hector, A., Inchausti, P., Lavorel, S.,&Wardle, D. A.(2005). Effects of biodiversity on ecosystem functioning:A consensus of current knowledge. Ecological Monographs, 75(1), 3-35.
Hossain, M. B.(2011). Macrobenthic community structure from a tropical estuary.Germany:LAP Lambert Academic Publishing GmbH&Co. Saarbrucken.
Hossain, M. B.,&Marshall, D. J.(2014). Benthic infaunal community structuring in an acidified tropical estuary. Aquatic Biosystems, 10, 11.
Hossain, M. B., Marshall, D. J.,&Venkatramanan, S.(2014). Sediment granulometry and organic matter content in the intertidal zone of the Sungai Brunei estuarine system, northwest coast of Borneo. Carpathian Journal of Earth and Environmental Sciences, 9(2), 231-239.
Koch, V.,&Wolff, M.(2002). Energy budget and ecological role of mangrove epibenthos in the Caete estuary. North Brazil. Marine Ecology Progress Series. 228,119-130.
Kroeker, K. J., Micheli, F., Gambi, M. C.,&Martz, T. R.(2011). Divergent ecosystem responses within a benthic marine community to Ocean acidification. PNAS,108, 14515-14520.
Ledger, M. E.,&Hildrew, A. G.(2005). The ecology of acidification and recovery:Changes in herbivore-algal food web linkages across a stream pH gradient.Environmental Pollution. 137(1), 103-118.
Li, S. W., Liu, Y. J., Li. F., Zhang, Y., Xu. Z. F., Lü, Z. B.,&Zhang, A. B.(2013). Macrobenthic functional groups in Laizhou Bay, east China. Chinese Journal of Ecology,32(2), 380-388(In Chinese).
Macdonald, T. A., Burd, B. J., Macdonald, V. I.,&van Roodselaar, A.(2010). Taxonomic and feeding guild classification for the marine benthic macroinvertebrates of the Strait of Georgia, British Columbia. Canadian Technical Report of Fisheries and Aquatic Sciences, 2874, 63.
Macdonald, T. A., Burd, B. J.,&van Roodselaar. A.(2012). Size structure of marine soft-bottom macrobenthic communities across natural habitat gradients:Implications for productivity and ecosystem function. PIOS One, 7(7), e40071.
Marshall, D. J., Santos, J. H., Leung, K. Y. M.,&Chak, W. H.(2008). Correlations between gastropod shell dissolution and water chemical properties in a tropical estuary. Marine Environmental Research, 66, 422-429.
Muniz, P.,&Pires, A. M. S.(1999). Trophic structure of polychaetes in the Sao Sebastiao Channel(Southern Brazil). Marine Biology, 134, 517-528.
Pearson, T.,&Rosenberg, R.(1978). Macrobenthic succession in relation to organic enrichment and pollution of the marine environment. Oceanography and Marine Biology:An Annual Review, 16, 229-311.
Pye, M. C., Vaughan, I. P.,&Ormerod, S. J.(2012). Episodic acidification affects the breakdown and invertebrate colonization of oak litter. Freshwater Biology, 57(11), 2318-2329.
Rhoads, D. C.,&Young, D. K.(1970). The Influence of deposit-feeding organisms on sediment stability and community trophic structure. Journal of Marine Research,28, 150-178.
Roach, A. C.(1997). The effect of acid water inflow on the estuarine benthic and fish communities in the Richmond River, NSW Australia. Australasian Journal of Ecotoxicology, 3, 25-56.
Rupert, E. E., Fox, R. S.,&Barnes, R. D.(2003). Invertebrate zoology:a functional evolutionary approach((7th Edition). California, USA:Saunders College Publishing.
Sahat, H. M. J. B.H.,&Howes, J.(1990). A study of the littoral ecosystem in Brunei estuary, Brunei Darussalam. The Brunei Museum Journal(pp. 100-164), 100-164.
Sivadas, S. K., Ingole. B., Ganesan, P., Sautya. S.,&Nanajkar, M.(2012). Role of environmental heterogeneity in structuring the macrobenthic community in a tropical sandy beach, west coast of India. Journal of Oceanography, 68(2),295-305.
Talley, T. S., Dayton, P. K.,&Ibarra-Obando, S. E.(2000). Tidal flat macrofaunal communities and their associated environments in estuaries of southern California and northern Baja California, Mexico. Estuaries, 23(1), 97-114.
Ter Braak, C. J. F.,&Prentice, I. C(1988). A theory of gradient analysis. Advances in Ecological Research, 18, 271-317.
Traister, E. M., McDowell, W. D., Kram, P., Fottova, D.,&Kolarikova, K.(2013). Persistent effects of acidification on stream ecosystem structure and function.Freshwater Science, 32, 586-596.
Weston, D. P.(1990). Quantitative examination of macrobenthic community changes along an organic enrichment gradient. Marine Ecology Progress Series,61, 233-244.
Xu, M. Z., Wang, Z. Y., Pan, B. Z.,&Zhao, N.(2012). Distribution and species composition of macroinvertebrates in the hyporheic zone of bed sediment. International Journal of Sediment Research, 27(2), 129-140.
Ysebaert, T., Herman, P. M. J., Meire, PR, Craeymeersch, J.,&Verbeek, H.(2003). Large scale spatial patterns in estuaries:Estuarine macrobenthic communities in the Schelde estuary. Estuarine, Coastal and Shelf Science, 57, 335-356.
Ysebaert, T., Meire, P., Coosen, J.,&Essink, K.(1998). Zonation of intertidal macrobenthos in the estuaries of Schelde and Ems. Aquatic Ecology, 32(1), 53-71.
Zhao, W. H., Wang, H. J., Wang, H. Z.,&Paul, G.(2011). Macroinvertebrates in the bed sediment of the Yellow River. International Journal of Sediment Research, 26(3), 255-268.
Amaral, V., Cabral, H. N.,&Bishop, M. J.(2014). Prior exposure influences the behavioural avoidance by an intertidal gastropod, Bembicium auratum, of acidified waters. Estuarine, Coastal and Shelf Science, 136, 82-90.
Bessa, F., Cunha, D., Concalves, S. C.,&Marques, J. C.(2013). Sandy beach macrofaunal assemblages as indicators of anthropogenic impacts on coastal dunes.Ecological Indicators, 30, 196-204.
Braeckman, U., Provoost, P., Gribsholt, B., van Gansbeke, D., Middelburg, J. J., Soetaert, K.,&Vanaverbeke, J.(2009). Role of macrofauna functional traits and density in biogeochemical fluxes and bioturbation. Marine Ecology Progress Series, 399, 173-186.
Brown, S. S., Gaston, G. R., Rakocinski, C. F..&Heard, R. W.(2000). Effects of sediment contaminants and environmental gradients on macrobenthic community trophic structure in Gulf of Mexico estuaries. Estuaries, 23(3), 411-424.
Carrasco, F. D.,&Carbajal, W.(1998). The distribution of polychaete feeding guilds in organic enriched sediments of San Vincente Bay, Central Chile. International Review of Hydrobiology,83,233-249.
Carstensen, J., Conley, D. J., Bonsdorff, E., Gustafsson, B. G., Hietanen, S., Janas, U.,&Voss, M.(2014). Hypoxia in the Baltic Sea:Biogeochemical cycles, benthic fauna, and management. Ambio, 43(1), 26-36.
Chardy, P.,&Clavier, J.(1988). Biomass and trophic structure of the macrobenthos in the south-west lagoon of New Caledonia. Marine Biology, 99, 105-202.
Clarke, K. R.(1993). Non-parametric multivariate analyses of changes in community structure. Australian Journal of Ecology, 18, 117-143.
Clarke, K. R.,&Warwick, R. M.(2001). Change in marine communities:An approach to statistical analysis and interpretation. Plymouth, UK:Plymouth Marine Laboratory.
Dauvin, J. C.,&Desroy, N.(2005). The food web in the lower part of the Seine estuary:A synthesis of existing knowledge. Hydrobiologia, 540(1-3), 13-27.
Dauwe, B., Herman, P. M. J.,&Heip, C. H. R.(1998). Community structure and bioturbation potential of macrofauna at four North Sea stations with contrasting food supply. Marine Ecology Progress Series, 173, 67-83.
Diaz, R. J.,&Schaffner, L C.(1990). The functional role of estuarine benthos In:M. Haired,&E. C. Krome(Eds.), Perspectives on the Chesapeake Bay, 1990Advances in Estuarine Sciences(pp. 25-56). U.S. Environmental Protection Agency, Chesapeake Bay Program(Report CBP/TRS/41/90).
Dimitriadis, C.,&Koutsoubas, D.(2011). Functional diversity and species turnover of benthic invertebrates along a local environmental gradient induced by an aquaculture unit:The contribution of species dispersal ability and rarity.Hydrobiologia, 670(1), 307-315.
Dove, C. M.,&Sammut,J.(2007). Impacts of estuarine acidification on survival and growth of Sydney rock oysters Saccostrea glomerata(Gould 1850).Journal of Shellfish Research, 26, 519-527.
Duan, X. H., Wang, Z. Y., Xu, M. Z.,&Zhang, K.(2009). Effect of streambed sediment on benthic ecology. International Journal of Sediment Research, 24(3), 325-338.
Fauchald, K.,&Jumars, P.(1979). The diet of worms:A study of polychaete feeding guilds. Oceanography and Marine Biology:An annual Review, 17,194-284.
Garcia-Arberas, L,&Rallo, A.(2002). The intertidal soft bottom infaunal macrobenthos in three Basque estuaries(Gulf of Biscay):A feeding guild approach.Hydrobiologia, 475-476, 457-468.
Gaston, G. R., Brown, S. S., Rakocinski, C. F., Heard, R. W.,&Summers, J. K.(1995).Trophic structure of macrobenthic communities in Northern Gulf of Mexico estuaries. Gulf Research Reports, 9,111-116.
Gaston, G. R., Rakocinski, C. F., Brown, S. S.,&Cleveland, C. M.(1998). Trophic function in estuaries:Response of macrobenthos to natural and contaminant gradients. Marine and Freshwater Research, 49(8), 833-846.
Gaudencio, M. J.,&Cabral, H. N.(2007). Trophic structure of macrobenthos in the Tagus estuary and adjacent coastal shelf. Hydrobiologia, 587(1), 241-251.
Gray, J. S.,&Elliott, M.(2009). Ecology of marine sediments:Science to management(2nd edition). Oxford, UK:Oxford University Press.
Grilo, T. F., Cardoso, P. G., Dolbeth, M., Bordalo, M. D.,&Pardal, M. A.(2011). Effects of extreme climate events on the macrobenthic communities'structure and functioning of a temperate estuary. Marine Pollution Bulletin, 62(2), 303-311.
Hammer, 0., Harper, D. A.,&Ryan, P. D.(2001). PAST:Paleontological statistics software package for education and data analysis. Palaeontologia Electronica, 4(1), 9.
Hanekom, N.(2011). Trophic structure and biomass distribution of macrobenthos on sheltered and semi-exposed rocky shores of Tsitsikamma Marine Protected Area. African Zoology, 46(2), 224-238.
Hooi, S. M. D.(1987). Sungai Brunei water quality and pollution study(Final Report).Brunei Darussalam:Government of Negara Brunei Darussalam.
Hooper, D. U., Chapin, F. S.,Ⅲ, Ewel, J. J., Hector, A., Inchausti, P., Lavorel, S.,&Wardle, D. A.(2005). Effects of biodiversity on ecosystem functioning:A consensus of current knowledge. Ecological Monographs, 75(1), 3-35.
Hossain, M. B.(2011). Macrobenthic community structure from a tropical estuary.Germany:LAP Lambert Academic Publishing GmbH&Co. Saarbrucken.
Hossain, M. B.,&Marshall, D. J.(2014). Benthic infaunal community structuring in an acidified tropical estuary. Aquatic Biosystems, 10, 11.
Hossain, M. B., Marshall, D. J.,&Venkatramanan, S.(2014). Sediment granulometry and organic matter content in the intertidal zone of the Sungai Brunei estuarine system, northwest coast of Borneo. Carpathian Journal of Earth and Environmental Sciences, 9(2), 231-239.
Koch, V.,&Wolff, M.(2002). Energy budget and ecological role of mangrove epibenthos in the Caete estuary. North Brazil. Marine Ecology Progress Series. 228,119-130.
Kroeker, K. J., Micheli, F., Gambi, M. C.,&Martz, T. R.(2011). Divergent ecosystem responses within a benthic marine community to Ocean acidification. PNAS,108, 14515-14520.
Ledger, M. E.,&Hildrew, A. G.(2005). The ecology of acidification and recovery:Changes in herbivore-algal food web linkages across a stream pH gradient.Environmental Pollution. 137(1), 103-118.
Li, S. W., Liu, Y. J., Li. F., Zhang, Y., Xu. Z. F., Lü, Z. B.,&Zhang, A. B.(2013). Macrobenthic functional groups in Laizhou Bay, east China. Chinese Journal of Ecology,32(2), 380-388(In Chinese).
Macdonald, T. A., Burd, B. J., Macdonald, V. I.,&van Roodselaar, A.(2010). Taxonomic and feeding guild classification for the marine benthic macroinvertebrates of the Strait of Georgia, British Columbia. Canadian Technical Report of Fisheries and Aquatic Sciences, 2874, 63.
Macdonald, T. A., Burd, B. J.,&van Roodselaar. A.(2012). Size structure of marine soft-bottom macrobenthic communities across natural habitat gradients:Implications for productivity and ecosystem function. PIOS One, 7(7), e40071.
Marshall, D. J., Santos, J. H., Leung, K. Y. M.,&Chak, W. H.(2008). Correlations between gastropod shell dissolution and water chemical properties in a tropical estuary. Marine Environmental Research, 66, 422-429.
Muniz, P.,&Pires, A. M. S.(1999). Trophic structure of polychaetes in the Sao Sebastiao Channel(Southern Brazil). Marine Biology, 134, 517-528.
Pearson, T.,&Rosenberg, R.(1978). Macrobenthic succession in relation to organic enrichment and pollution of the marine environment. Oceanography and Marine Biology:An Annual Review, 16, 229-311.
Pye, M. C., Vaughan, I. P.,&Ormerod, S. J.(2012). Episodic acidification affects the breakdown and invertebrate colonization of oak litter. Freshwater Biology, 57(11), 2318-2329.
Rhoads, D. C.,&Young, D. K.(1970). The Influence of deposit-feeding organisms on sediment stability and community trophic structure. Journal of Marine Research,28, 150-178.
Roach, A. C.(1997). The effect of acid water inflow on the estuarine benthic and fish communities in the Richmond River, NSW Australia. Australasian Journal of Ecotoxicology, 3, 25-56.
Rupert, E. E., Fox, R. S.,&Barnes, R. D.(2003). Invertebrate zoology:a functional evolutionary approach((7th Edition). California, USA:Saunders College Publishing.
Sahat, H. M. J. B.H.,&Howes, J.(1990). A study of the littoral ecosystem in Brunei estuary, Brunei Darussalam. The Brunei Museum Journal(pp. 100-164), 100-164.
Sivadas, S. K., Ingole. B., Ganesan, P., Sautya. S.,&Nanajkar, M.(2012). Role of environmental heterogeneity in structuring the macrobenthic community in a tropical sandy beach, west coast of India. Journal of Oceanography, 68(2),295-305.
Talley, T. S., Dayton, P. K.,&Ibarra-Obando, S. E.(2000). Tidal flat macrofaunal communities and their associated environments in estuaries of southern California and northern Baja California, Mexico. Estuaries, 23(1), 97-114.
Ter Braak, C. J. F.,&Prentice, I. C(1988). A theory of gradient analysis. Advances in Ecological Research, 18, 271-317.
Traister, E. M., McDowell, W. D., Kram, P., Fottova, D.,&Kolarikova, K.(2013). Persistent effects of acidification on stream ecosystem structure and function.Freshwater Science, 32, 586-596.
Weston, D. P.(1990). Quantitative examination of macrobenthic community changes along an organic enrichment gradient. Marine Ecology Progress Series,61, 233-244.
Xu, M. Z., Wang, Z. Y., Pan, B. Z.,&Zhao, N.(2012). Distribution and species composition of macroinvertebrates in the hyporheic zone of bed sediment. International Journal of Sediment Research, 27(2), 129-140.
Ysebaert, T., Herman, P. M. J., Meire, PR, Craeymeersch, J.,&Verbeek, H.(2003). Large scale spatial patterns in estuaries:Estuarine macrobenthic communities in the Schelde estuary. Estuarine, Coastal and Shelf Science, 57, 335-356.
Ysebaert, T., Meire, P., Coosen, J.,&Essink, K.(1998). Zonation of intertidal macrobenthos in the estuaries of Schelde and Ems. Aquatic Ecology, 32(1), 53-71.
Zhao, W. H., Wang, H. J., Wang, H. Z.,&Paul, G.(2011). Macroinvertebrates in the bed sediment of the Yellow River. International Journal of Sediment Research, 26(3), 255-268.