Macrozoobenthos in Yangtze Floodplain Lakes
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Macrozoobenthos in Yangtze floodplain lakes: patterns of density, biomass, and production in relation to river connectivity Author(s): Bao-Zhu Pan, Hai-Jun Wang, Xiao-Min Liang, and Hong-Zhu Wang Source: Journal of the North American Benthological Society, 30(2):589-602. 2011. Published By: North American Benthological Society DOI: 10.1899/10-025.1 URL: http://www.bioone.org/doi/full/10.1899/10-025.1 BioOne (www.bioone.org) is an electronic aggregator of bioscience research content, and the online home to over 160 journals and books published by not-for-profit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. 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Soc., 2011, 30(2):589–602 ’ 2011 by The North American Benthological Society DOI: 10.1899/10-025.1 Published online: 5 April 2011 Macrozoobenthos in Yangtze floodplain lakes: patterns of density, biomass, and production in relation to river connectivity 1 2 3 4 Bao-Zhu Pan , Hai-Jun Wang , Xiao-Min Liang , AND Hong-Zhu Wang State Key Laboratory of Freshwater Ecology and Biotechnology, Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan, Hubei 430072, China Abstract. A systematic investigation of macrozoobenthos was conducted in Yangtze floodplain waters to reveal patterns of density, biomass, and production in relation to river connectivity. In the Yangtze- connected lakes, 78 taxa belonging to 33 families and 62 genera were identified. Macrozoobenthos density was 327 individuals/m2, biomass was 1.40 g dry mass/m2, and production was 3.23 g dry mass m22 y21. The assemblages were characterized by high diversity, high production, and high bivalve-filterer abundance. The key factor determining the macrozoobenthic assemblages was river connectivity. As river connectivity increased, 3 types of response patterns were observed: 1) density, biomass, and production of collector-filterers (mainly Bivalvia), shredders (e.g., Stictochironomus), and predators (e.g., Dytiscidae) showed unimodal changes, i.e., first increased and then decreased; 2) density, biomass, and production of collector-gatherers (mainly Tubificidae and Chironomidae) decreased continuously; and 3) density of scrapers (mainly Gastropoda) decreased, whereas their biomass and production changed unimodally. At an intermediate level of river connectivity, biomass and production of total macrozoobenthos reached maxima, whereas density decreased with increasing river connectivity. Previous research showed that a diversity of zoobenthos also peaks at moderate connectivity with rivers. Therefore, to maintain high productivity as well as high biodiversity in the Yangtze floodplain, protecting the remnants of river- connected lakes and linking disconnected lakes freely with the mainstream are crucial. Key words: macrozoobenthos, water current, connectivity, Yangtze floodplain lakes. The riverine floodplain is one of the most endan- are key factors structuring macrozoobenthic assem- gered landscapes worldwide (Tockner and Stanford blages (Van den Brink and Van der Velde 1991, 2002). In floodplain waters, macrozoobenthos play an Amoros and Bornette 2002, Paillex et al. 2007). important role in trophic dynamics by cycling Alpha diversity of macrozoobenthos reaches a nutrients and providing food for higher trophic levels, maximum at an intermediate level of river–floodplain such as fish and birds (Van den Brink et al. 1994). A connectivity (Obrdlik and Fuchs 1991, Ward 1998, better understanding of macrozoobenthic assemblag- Tockner et al. 1999b, Ward et al. 1999, Amoros and es is of great significance to floodplain conservation Bornette 2002), whereas b diversity tends to be higher and management. when river connectivity is weaker (Castella et al. 1991, Previous research on macrozoobenthos in flood- Reckendorfer et al. 2006). The relationship between plains has been mostly restricted to the Danube, macrozoobenthic standing crops and hydroperiod has Rhine, and Mississippi basins (Castella et al. 1991, been studied less often (Corti et al. 1997, Flinn et al. Obrdlik and Fuchs 1991, Van den Brink and Van der 2008), but knowledge of responses of macrozoo- Velde 1991, Van den Brink et al. 1994, Tockner et al. benthic standing crops and production to river 1999b). Hydrological connectivity with rivers and connectivity is important to ecosystem management. pertinent variables, i.e., flood-pulse duration, flood Therefore, our goal was to explore the patterns of frequency, water velocity, and water-level fluctuation, standing crops and production of zoobenthos along river connectivity gradients. 1 E-mail addresses: [email protected] In the Yangtze basin of China, floodplain lakes are 2 [email protected] numerous, with a total area .16,600 km2. Historically, 3 [email protected] most lakes were connected freely with the main river 4 To whom correspondence should be addressed. E-mail: course of the Yangtze, where floods occur periodical- [email protected] ly. To protect villages and cultivated land along the 589 590 B. PAN ET AL. [Volume 30 lakeshore from flooding, embankments and sluice same habitat adjacent to the benthic sampling site, gates were constructed between the 1950s and the then cleaned and weighed for determination of wet 1970s. These structures eventually isolated most lakes mass (BMac). Sediment samples of 500 mL were taken from the river. At present, only 3 medium- to large- from the hyporheic zone (depth: 0–0.15 m), and sized lakes (Dongting Lake, Poyang Lake, and Shijiu particle size was analyzed using the sieving method. Lake) remain connected with the Yangtze main- Sediment was classified according to the Wentworth stream. Although they are now few in number, these scale. Particle diameters of coarse sand, medium sand, river-connected lakes are important for maintaining fine sand, silt, and clay were 0.5 to 2, 0.25 to 0.5, 0.0625 the unique and diverse biota of the entire Yangtze to 0.25, 0.004 to 0.0625, and ,0.004 mm, respectively ecosystem. However, our knowledge concerning the (Allan and Castillo 2007). macrozoobenthos of these lakes is still limited, and Quantitative samples of macrozoobenthos were only a few works describing these assemblages have taken from the hyporheic zones with a weighted been published (Dai et al. 2000, Wang et al. 2007, Xie Petersen grab (0.0625 m2 3 0.15 m) and then passed et al. 2007). through a 420-mm sieve. Specimens were sorted Our paper reports the results of a systematic manually from sediment on a white porcelain plate investigation of macrozoobenthos in all 3 Yangtze- and preserved in 10% formalin. Wet mass of connected lakes. The purpose of our study was 2-fold: zoobenthos was determined with an electronic bal- 1) to describe the overall character of macrozoo- ance after the specimens were blotted, and dry mass benthic assemblages in these lakes, and 2) to reveal (mollusks without shells) was calculated according to patterns of density, biomass, and production of the ratios of dry:wet mass and tissue-shell weight different macrozoobenthic groups in relation to river reported by Yan and Liang (1999). In the study by Yan connectivity. and Liang (1999), the data were collected from mid- lower Yangtze lakes, and mollusk samples were Methods treated according to the method described by Banse and Mosher (1980) (the shells were removed and then Study area shell and tissue were dried separately). Production of Dongting Lake, Poyang Lake, and Shijiu Lake are zoobenthos was calculated mainly according to the situated in mid-to-lower Yangtze basin in the mon- ratio of production to biomass (P/B) reported from soon region of the East Asia subtropical zone. Yangtze lakes (Yan 1998). In the study by Yan (1998), Locations, sampling sites, and limnological variables 12 samples of macrozoobenthos were taken monthly of study lakes are given in Fig. 1 and Table 1. over a period of 1 y from 2 lakes, and the production Field investigations were conducted in May to July was estimated by the size–frequency method. The (high-water level) and September to December (low- P/B ratio of Corbicula was taken from Marsh (1985). For water level) 2004. The data from the 2 investigations taxa without production data, P/B ratios of similar were combined for analyses. Water depth (Z) and forms were used (Liang and Wang 1999). All taxa were Secchi depth (ZSD) were measured with a sounding assigned to functional feeding groups (shredders, lead and a Secchi disc, respectively. Water velocity collector-gatherers, collector-filterers, scrapers, and (U) was measured with a propeller-type current meter predators) according to Morse et al. (1994) and Liang (Model LS 1206B; Nanjing Automation Institute of and Wang (1999). When a taxon had several possible Water Conservancy and Hydrology, Nanjing, China). feeding activities, its functional designations were One-liter