Natural Leaf Packs and Invertebrates in a Tropical Stream: Is There a Functional Relationship?
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Sri Lanka J. Aquat. Sci. 24(2) (2019): 65-76 http://doi.org/10.4038/sljas.v24i2.7568 Natural leaf packs and invertebrates in a tropical stream: Is there a functional relationship? Hiranthi Walpola1, Maria Leichtfried2 and Leopold Füreder1 1River Ecology and Conservation, Institute of Ecology, University of Innsbruck, Technikerstr. 25, A-6020 Innsbruck, Austria 2Institute for Limnology of the Austrian Academy of Sciences, Mondseestr. 9, A-5310 Mondsee, Austria *Correspondence ([email protected]) https://orcid.org/0000-0002-5793-7270 Abstract Naturally occurring leaf packs in five sites along the elevation gradient in Eswathu Oya stream in Sri Lanka were sampled and analysed for species composition of leaves and benthic macroinvertebrates. Eswathu Oya is a perennial, low order, mountain stream in the wet climatic zone of Sri Lanka. Leaves of only five species of plants were found in all sites – Hevea brasiliensis, Ochlandra stridula, Alstonia macrophylla, Terminalia arjuna and Melia azedarach. In each sampling site Hevea brasiliensis and Ochlandra stridula leaves were found in higher abundance. The contribution of other leaves varied with the site. Diptera larvae of the family Chironomidae, were the dominant group of leaf colonizing animals. The relative abundance of Chironomidae decreased along the elevation gradient in the down-stream. Gathering Collectors (GC) were the dominant functional feeding group (FFG) found in all leaf packs, but their abundance decreased along the downstream elevation gradient. Shredders were not present, as reported from some other tropical streams. Predators and Scrapers were increased along the downstream gradient. Filtering Collectors and Pierce-Herbivores did not show a clear pattern. The study documents that macro invertebrate communities responded to change in water quality, availability of leaves and their nutritional quality along the elevation gradient in Eswathu Oya, Sri Lanka. Keywords: Lotic ecosystems, Longitudinal gradient, Functional feeding groups, Tropical streams INTRODUCTION considered as the dominant determinants of the decomposition process of allochthonous organic Allochthonous input of organic matter as a matter in small temperate streams Dobson et al., source of nutrients can influence the structure and 2002). However, the role of invertebrate organisms function of food webs in streams (Polis and Hurd, in the decomposition has not been satisfactorily 1996; Anderson and Polis, 1998; Leichtfried, investigated in tropical streams (Dobson et al., 2007). Different physical, chemical and biological 2002; Mathuriau and Chauver, 2002). processes in the stream contribute to the Decomposition processes in natural decomposition of organic matter, mineralizing ecosystems are also influenced by differences in them to elements, which are released to the stream temperature, composition of the organic matter, to be available for micro-organisms and biofilms and structure of decomposer assemblages (Swift et (Swift et al. 1979; Gessnnet et al., 1999). These al., 1979; Aerts, 1997; Royer and Minshall, 2003; processes can increase the production of primary Hättenschwiler and Gasser, 2005). Litter consumers; thus creating more food for higher consumption by invertebrates may fluctuate in trophic levels (Strong, 1992; Huxel and McCann, response to litter availability and palatability 1998). Feeding activities and decomposition (Cummins et al., 1989), leaf structure and stimulated by these invertebrates vary with time chemistry (Campbell and Fuchshuber, 1995; and space. Invertebrates are important in the Findlay et al., 1996), microbial conditioning, decomposition process, through accelerating it by fragmenting the organic matter into smaller pieces, thus increasing the surface area available for biofilms and microbial activity (Graça, 2001; Leichtfried, 2007). Shredders were often 66 Hiranthi Walpola et al. development of biofilms (Bärlocher and Kendrick, factors like the local water flow regime (e.g., 1975) and other parameters. Brown and Brussock, 1991), substrate composition Some studies report that in streams, the and porosity (Wood and Armitage, 1997), substrate composition of macro invertebrate assemblages stability (Death, 1995), and the presence of large colonizing leaf bags differs between tropical and wood (Benke et al., 1984) play an important role in temperate systems, with shredders as the dominant structuring benthic communities, but group in decomposing leaf litter in temperate comparatively little information on these subjects regions (Vannote et al., 1980; Graça, 2001). In is available from tropical areas. tropical regions, shredders are often not the Very little attention has been given to study dominant colonizing group (Rosemond et al., invertebrates associated with the leaf litter 1989; Dudgeon and Wu, 1999; Dudgeon, 2000; decomposition process in tropical countries, Dobson et al., 2002), but some other studies report especially in South East Asia, in contrast to Hong the opposite (e.g., Pearson et al., 1989; Cheshire et Kong, where several studies are published (e.g., Li al., 2005; Cummins et al., 2005). and Dudgeon, 2009). Most studies of the benthic Variations in aquatic habitats such as streams fauna in Sri Lanka´s streams were taxonomic or have been reviewed by Webster and Benfield faunistic (e.g., Wijenayake et al., 2001). (1986) and Royer and Minshall (2003), who The aim of this study was (i) to evaluate the included differences due to physical and chemical natural density and type of senescent leaves in a factors (Chergui and Pattèe, 1988; Irons et al., tropical stream along a longitudinal gradient, (ii) to 1994; Subrkropp and Chauvet, 1995; Dangles et estimate potential habitat and food qualities for al., 2004), type of riparian forest and land use macro invertebrates in their natural environment, patterns (Whiles and Wallace, 1997; Sponseller and (iii) to analyse potential functional and Benfield, 2001; Danger and Robson, 2004), relationships between leaves and invertebrates. and the effects of litter quality (Petersen and Cummins, 1974; Webster and Benfield, 1986) and MATERIALS AND METHODS macro invertebrates (Wallace and Webster, 1996; Rosmend et al., 1998; Graça, 2001; Wright and Study area Covich, 2005). Large differences were found among experimental treatments, where macro The investigation was carried out in the Eswathu invertebrate abundance was manipulated (Graça, Oya, a low order stream in the wet climatic zone of 2001), with some studies showing higher Sri Lanka (Fig. 1). The river drains a hilly, decomposition rates, when macro invertebrates cascading landscape and runs into the Kelani were present (Petersen and Cummins, 1974; River, the second largest river (144 km long) in Sri Bradford et al., 2002). Other studies did not show Lanka. The catchment is geologically dominated any significant effects due to macro invertebrates by the charnockite series in this area. Due to the (Rosemond et al., 1998; Stockley et al., 1998) or topology of the catchment and the fluctuation of an interaction with leaf quality (Vanconcelos and water level, pools and riffles are developed in Laurance, 2005; Wright and Covich, 2005). Eswathu Oya stream, which provide the basic Therefore, leaf litter decomposition processes are niche structure for the benthic invertebrates. strongly influenced by differences in climatic Eswathu Oya has large consecutive pools and conditions, habitat types, and by effects of riffles through the natural cascade in the landscape. different functional feeding groups (FFGs) of Five sites along the longitudinal gradient were decomposer assemblages. selected for sampling. The temperature, pH, Macro invertebrate community structure and electrical conductivity, water velocity and profiles function is influenced by a complex of abiotic and were recorded in order to calculate water discharge biotic factors that interact over a range of spatial in pools in each site. and temporal scales (Richards et al., 1997). Abiotic 67 Hiranthi Walpola et al. Fig 1 Study site locations (1 – 5) in Eswathu Oya Sampling method Five random samples of natural leaf packs (fallen/ stream. The temperature showed slight increment, senescent leaves in the bottom of the stream) were the dissolved oxygen and electrical conductivity collected from each site using the Dip net method shows also a slight increment towards downstream. (100 µm, 20 × 20 cm2 sampling area). The species The slightly acidic pH remains throughout the composition of the plant leaves in the natural leaf sites. The discharge increased 15 times from packs was determined in the lab and benthic macro headwaters to downstream (Table 1). invertebrates were separated and identified up to Altogether, 29 species of plant leaves were the best possible taxonomic level. The benthic found in the leaf packs. Leaves of plant species macro invertebrates were classified into functional such as Hevea brasilensis, Ochlandra stridula, feeding groups (FFGs) according to Merritt and Alstonia macrophylla, Terminalia arjuna, Melia Cummins (1996), Dudgeon (1999) and Fernando azedarach and Artocarpus nobilis occurred in all and Weerawardhane (2002). sampling site. Ochlandra stridula and Hevea brasiliensis were representing as the most RESULTS dominant plant leaves in leaf packs and found in head waters to downstream sites (Table 2). The A total of 25 natural leaf packs (in total 500 leaves) mean quantitative occurrence of different species were analyzed from five sites along the in the leaf packs is shown in Table 3\ longitudinal gradient (Fig. 1) of Eswathu Oya 68 Hiranthi Walpola et al. Table 1 Surface water characteristics of the investigated five sites in longitudinal profile of Eswathu Oya, measured during sampling (mean±SD, n = 5). Sites Parameter 1 2 3 4 5 Temp. (°C) 25.6±0.6 25.9±0.8 26.2±0.3 26.4±0.2 26.8±0.4 pH 6.1±0.2 6.1±0.1 6.2±0.4 6.1±0.5 5.9±0.8 Dissolve Oxygen (mg l-1) 6.4±0.1 6.8±0.6 6.9±0.4 6.9±0.3 7.1±0.5 Discharge (l sec-1) 13.06±5.2 30.93±7.3 65.49±5.9 85.49±89 209.12±10.5 -1 Conductivity (µS20 cm ) 29.2±0.1 31.2±0.3 32.0±0.2 34.2±0.2 33.2±0.1 Table 2 Leaf species composition of sampled leaf packs in Eswathu Oya.