The functional role of burrowing bivalves in freshwater ecosystems

1. Freshwater systems are losing biodiversity at a rapid rate, yet we know little about the functional role of most of this biodiversity. The ecosystem roles of freshwater burrowing bivalves have been particularly understudied. Here we summarize what is known about the functional role of burrowing bivalves in the orders Unionoida and Veneroida in lakes and streams globally. 2. Bivalves filter phytoplankton, bacteria and particulate organic matter from the water column. Corbicula and sphaeriids also remove organic matter from the sediment by deposit feeding, as may some unionids. Filtration rate varies with bivalve species and size, temperature, particle size and concentration, and flow regime. 3. Bivalves affect nutrient dynamics in freshwater systems, through excretion as well as biodeposition of faeces and pseudofaeces. Excretion rates are both size and species dependent, are influenced by reproductive stage, and vary greatly with temperature and food availability. 4. Bioturbation of sediments through bivalve movements increases sediment water and oxygen content and releases nutrients from the sediment to the water column. The physical presence of bivalve shells creates habitat for epiphytic and epizoic organisms, and stabilizes sediment and provides refugia for benthic fauna. Biodeposition of faeces and pseudofaeces can alter the composition of benthic communities. 5. There is conflicting evidence concerning the role of resource limitation in structuring bivalve communities. Control by bivalves of primary production is most likely when their biomass is large relative to the water volume and where hydrologic residence time is long. Future studies should consider exactly what bivalves feed upon, whether feeding varies seasonally and with habitat, and whether significant overlap in diet occurs. In particular, we need a clearer picture of the importance of suspension versus deposit feeding and the potential advantages and tradeoffs between these two feeding modes. 6. In North America, native burrowing bivalves (Unionidae) are declining at a catastrophic rate. This significant loss of benthic biomass, coupled with the invasion of an exotic burrowing bivalve (Corbicula), may result in large alterations of ecosystem processes and functions.     

Community Biomass and Bottom up Multivariate Nutrient Complementarity Mediate the Effects of Bioturbator Diversity on Pelagic Production

Tests of the biodiversity and ecosystem functioning (BEF) relationship have focused little attention on the importance of interactions between species diversity and other attributes of ecological communities such as community biomass. Moreover, BEF research has been mainly derived from studies measuring a single ecosystem process that often represents resource consumption within a given habitat. Focus on single processes has prevented us from exploring the characteristics of ecosystem processes that can be critical in helping us to identify how novel pathways throughout BEF mechanisms may operate. Here, we investigated whether and how the effects of biodiversity mediated by non-trophic interactions among benthic bioturbator species vary according to community biomass and ecosystem processes. We hypothesized that (1) bioturbator biomass and species richness interact to affect the rates of benthic nutrient regeneration [dissolved inorganic nitrogen (DIN) and total dissolved phosphorus (TDP)] and consequently bacterioplankton production (BP) and that (2) the complementarity effects of diversity will be stronger on BP than on nutrient regeneration because the former represents a more integrative process that can be mediated by multivariate nutrient complementarity. We show that the effects of bioturbator diversity on nutrient regeneration increased BP via multivariate nutrient complementarity. Consistent with our prediction, the complementarity effects were significantly stronger on BP than on DIN and TDP. The effects of the biomass-species richness interaction on complementarity varied among the individual processes, but the aggregated measures of complementarity over all ecosystem processes were significantly higher at the highest community biomass level. Our results suggest that the complementarity effects of biodiversity can be stronger on more integrative ecosystem processes, which integrate subsidiary “simpler” processes, via multivariate complementarity. In addition, reductions in community biomass may decrease the strength of interspecific interactions so that the enhanced effects of biodiversity on ecosystem processes can disappear well before species become extinct.     

Weak response of saltmarsh infauna to ecosystem-wide nutrient enrichment and fish predator reduction: A four-year study

We examined the effect of whole-ecosystem manipulations of predator removal and nutrient enrichment on saltmarsh macroinfauna in the Plum Island Estuary, Massachusetts. Nitrate and phosphate loading rates were increased 10× above background levels in experimental creeks, and we significantly reduced (by 60%) the abundance of the killifish, Fundulus heteroclitus, a key predator in this system. Two creek pairs were manipulated; Creek Pair 1 for three growing seasons and Creek Pair 2 for one. Infaunal responses were examined in four habitats along the inundation gradient: mudflat, creek wall, Spartina alterniflora, and S. patens habitats. Although benthic microalgae increased synergistically in our treatments, we detected no long-term, population-level numerical response by any taxon. Similarly, no long-term species diversity or community responses were observed. However, nutrient enrichment increased the population biomass of the polychaete Manayunkia aestuarina in the creek wall habitat and the oligochaete Cernosvitoviella immota in the S. alterniflora habitat. No numerical or biomass responses of infauna were detected in predator removal treatments although indirect effects associated with killifish reduction may have contributed to an ephemeral interaction between nutrient addition and predator reduction in S. patens habitat. Our data suggest that population and community responses between benthic microalgae and macroinfauna are not tightly coupled even though some species benefit from increased benthic algae biomass by achieving larger body size.     

Effects of benthivorous fish on biogeochemical processes in lake sediments

1. Studies of aquatic environments have shown that community organisation may strongly affect ecosystem functioning. One common phenomenon is a change in nutrient level following a shift in the fish community composition. Although several hypotheses have been suggested, there is no consensus on which mechanisms are involved. Our study evaluated indirect effects of benthivorous fish on the biogeochemical processes at the sediment–water interface separately from direct effects caused by nutrient excretion or sediment resuspension. 2. We assigned field enclosures to three treatments representing typical pond communities; without fish, addition of approximately 10 small tench or addition of one large bream. After one summer, we monitored the water chemistry, benthic invertebrates and periphyton in the enclosures and sampled sediment cores for laboratory analysis of biochemical process rates (oxygen, phosphorus and nitrogen exchange between sediment and water, and denitrification rate). 3. Fish had strong negative effects on benthic invertebrates, but weaker effects on periphyton, organic content and porosity of the sediment. Moreover, there were significant positive fish effects on both phosphorus and nitrogen concentrations in the water. However, there were no general treatment effects on sediment processes that could explain the treatment effects on water chemistry in the enclosures. 4. Hence, overall treatment effects attenuated along the chain of interactions. We conclude that the observed effect of benthic fish on water chemistry was probably because of direct effects on nutrient excretion or resuspension of sediment. The similarity between bream and tench treatments suggests large niche complementarity despite their different habitat preferences.     

REGULAR PAPERS Habitat distribution and size structure in freshwater fish communities: effects of vendace on interactions between perch and roach

In a study of habitat selection and size structure in three fish species in 115 Swedish lakes, roach Rutilus rutilus used the pelagic habitat to a lesser extent and were lower in pelagic biomass in the presence of vendace Coregonus albula. Size distributions of roach were skewed toward larger sizes in lakes with vendace, although a similar pattern of size-dependent habitat use was found regardless of lake group. In disagreement with predictions, vendace did not affect relative perch Perca fluviatilis biomass, size structure, or the proportion of large perch. However, a higher growth for large perch indicated higher per capita energy intake in lakes with vendace.     

Effects of chronic bottom trawling disturbance on benthic biomass, production and size spectra in different habitats

Bottom trawling has widespread impacts on benthic communities and habitats. It is argued that the impact of chronic bottom trawling on benthic infauna depends on the natural disturbance levels to which benthic communities are adapted. We analysed biomass, production and size structure of two communities from a muddy and a sandy habitat, in relation to quantified gradients of trawling disturbance on real fishing grounds. We used an allometric relationship between body mass and individual production to biomass ratio to estimate community production. Chronic trawling had a negative impact on the biomass and production of benthic communities in the muddy habitat, while no impact was identified on benthic communities from the sandy habitat. These differences are the result of differences in size structure within the two communities that occur in response to increasing trawling disturbance.     

Effects of retrogressive permafrost thaw slumping on sediment chemistry and submerged macrophytes in Arctic tundra lakes

1. Global warming is predicted to cause changes in permafrost cover and stability in the Arctic. Zones of high ion concentration in regions of ice‐rich permafrost are a reservoir of chemicals that can potentially be transferred to fresh waters during thawing. Consequently, input of enriched runoff from the thaw and sediment and vegetation from the landscape could alter lakes by affecting their geochemistry and biological production. 2. Three undisturbed lakes and five lakes disturbed by retrogressive permafrost thaw slumps were sampled during late summer of 2006 to assess the potential effects of thermokarst shoreline slumping on water and sediment chemistry, the underwater light regime, and benthic macrophyte biomass and community structure. 3. Undisturbed lakes had sediments rich in organic material and selected micronutrients, while disturbed lakes had sediments richer in calcium, magnesium and strontium, greater transparency of the water column, and a well‐developed submerged macrophyte community. 4. It is postulated that enriched runoff chemistry may alter nutrient availability at the sediment–water interface and also the degradation of organic material, thus affecting lake transparency and submerged macrophytes. The results suggest that retrogressive permafrost slumping can significantly affect food webs in arctic tundra lakes through an increase in macrophyte biomass and development of a more complex benthic habitat.     

Pacific walrus: Benthic bioturbator of Beringia

The dependency of walruses on sea ice as habitat, the extent of their feeding, their benthic bioturbation and consequent nutrient flux suggest that walruses play a major ecological role in Beringia. This suggestion is supported by several lines of evidence, accumulated during more than three decades of enquiry and leading to the hypothesis that positive feedbacks of walrus feeding strongly influence productivity and ecological function via benthic bioturbation and nutrient flux. Walruses annually consume an estimated 3 million metric tons of benthic biomass. Walrus prey species inhabit patches across the shelf according to sediment type and structure. Side-scan sonar and our calculations indicate that the area affected by walrus feeding is in the order of thousands of square kilometers per year. Annual to long-term walrus bioturbation results in significant, large-scale changes in sediment and biological-community structure, and magnifies nutrient flux from sediment pore water to the water column by about two orders of magnitude over wide areas. The combined effects of walrus feeding must be placed in the context of long-term, regional climate changes and responses. Should sea ice continue to move northward as a result of climate change, the walrus' ecological role could be diminished or lost, the benthic ecosystem could be fundamentally altered and native subsistence hunters would be deprived of important resources.     

The effects of storms,heavy rain,and sedimentation on the shallow coral reefs of St. John,US Virgin Islands

Efforts to limit plant growth in streams by reducing nutrients would benefit from an understanding of the relative influences of nutrients, streamflow, light, and other potentially important factors. We measured macrophytes, benthic algae, nutrients in water and sediment, discharge, and shading from 30 spring-fed or runoff-influenced streams in the upper Snake River basin, ID, USA. We hypothesized that in hydrologically stable, spring-fed streams with clear water, macrophyte and benthic algae biomass would be a function of bioavailable nutrients in water or sediments, whereas in hydrologically dynamic, runoff-influenced streams, macrophyte and benthic algae biomass would further be constrained by flow disturbance and light. These hypotheses were only partly supported. Nitrogen, both in sediment and water, was positively correlated with macrophyte biomass, as was loosely sorbed phosphorus (P) in sediment. However, P in water was not. Factors other than nutrient enrichment had the strongest influences on macrophyte species composition. Benthic algal biomass was positively correlated with loosely sorbed sediment P, lack of shade, antecedent water temperatures, and bicarbonate. These findings support the measurement of bioavailable P fractions in sediment and flow histories in streams, but caution against relying on macrophyte species composition or P in water in nutrient management strategies for macrophytes in streams.     

Consequences of Increasing Hypoxic Disturbance on Benthic Communities and Ecosystem Functioning

Disturbance-mediated species loss has prompted research considering how ecosystem functions are changed when biota is impaired. However, there is still limited empirical evidence from natural environments evaluating the direct and indirect (i.e. via biota) effects of disturbance on ecosystem functioning. Oxygen deficiency is a widespread threat to coastal and estuarine communities. While the negative impacts of hypoxia on benthic communities are well known, few studies have assessed in situ how benthic communities subjected to different degrees of hypoxic stress alter their contribution to ecosystem functioning. We studied changes in sediment ecosystem function (i.e. oxygen and nutrient fluxes across the sediment water-interface) by artificially inducing hypoxia of different durations (0, 3, 7 and 48 days) in a subtidal sandy habitat. Benthic chamber incubations were used for measuring responses in sediment oxygen and nutrient fluxes. Changes in benthic species richness, structure and traits were quantified, while stress-induced behavioral changes were documented by observing bivalve reburial rates. The initial change in faunal behavior was followed by non-linear degradation in benthic parameters (abundance, biomass, bioturbation potential), gradually impairing the structural and functional composition of the benthic community. In terms of ecosystem function, the increasing duration of hypoxia altered sediment oxygen consumption and enhanced sediment effluxes of NH₄ ⁺ and dissolved Si. Although effluxes of PO₄ ³⁻ were not altered significantly, changes were observed in sediment PO₄ ³⁻ sorption capability. The duration of hypoxia (i.e. number of days of stress) explained a minor part of the changes in ecosystem function. Instead, the benthic community and disturbance-driven changes within the benthos explained a larger proportion of the variability in sediment oxygen- and nutrient fluxes. Our results emphasize that the level of stress to the benthic habitat matters, and that the link between biodiversity and ecosystem function is likely to be affected by a range of factors in complex, natural environments.     

Climate warming and agricultural stressors interact to determine stream macroinvertebrate community dynamics

Global climate change is likely to modify the ecological consequences of currently acting stressors, but potentially important interactions between climate warming and land‐use related stressors remain largely unknown. Agriculture affects streams and rivers worldwide, including via nutrient enrichment and increased fine sediment input. We manipulated nutrients (simulating agricultural run‐off) and deposited fine sediment (simulating agricultural erosion) (two levels each) and water temperature (eight levels, 0–6°C above ambient) simultaneously in 128 streamside mesocosms to determine the individual and combined effects of the three stressors on macroinvertebrate community dynamics (community composition and body size structure of benthic, drift and insect emergence assemblages). All three stressors had pervasive individual effects, but in combination often produced additive or antagonistic outcomes. Changes in benthic community composition showed a complex interplay among habitat quality (with or without sediment), resource availability (with or without nutrient enrichment) and the behavioural/physiological tendency to drift or emerge as temperature rose. The presence of sediment and raised temperature both resulted in a community of smaller organisms. Deposited fine sediment strongly increased the propensity to drift. Stressor effects were most prominent in the benthic assemblage, frequently reflected by opposite patterns in individuals quitting the benthos (in terms of their propensity to drift or emerge). Of particular importance is that community measures of stream health routinely used around the world (taxon richness, EPT richness and diversity) all showed complex three‐way interactions, with either a consistently stronger temperature response or a reversal of its direction when one or both agricultural stressors were also in operation. The negative effects of added fine sediment, which were often stronger at raised temperatures, suggest that streams already impacted by high sediment loads may be further degraded under a warming climate. However, the degree to which this will occur may also depend on in‐stream nutrient conditions.     

Bottom‐up and top‐down effects of browning and warming on shallow lake food webs

Productivity and trophic structure of aquatic ecosystems result from a complex interplay of bottom‐up and top‐down forces that operate across benthic and pelagic food web compartments. Projected global changes urge the question how this interplay will be affected by browning (increasing input of terrestrial dissolved organic matter), nutrient enrichment and warming. We explored this with a process‐based model of a shallow lake food web consisting of benthic and pelagic components (abiotic resources, primary producers, grazers, carnivores), and compared model expectations with the results of a browning and warming experiment in nutrient‐poor ponds harboring a boreal lake community. Under low nutrient conditions, the model makes three major predictions. (a) Browning reduces light and increases nutrient supply; this decreases benthic and increases pelagic production, gradually shifting productivity from the benthic to the pelagic habitat. (b) Because of active habitat choice, fish exert top‐down control on grazers and benefit primary producers primarily in the more productive of the two habitats. (c) Warming relaxes top‐down control of grazers by fish and decreases primary producer biomass, but effects of warming are generally small compared to effects of browning and nutrient supply. Experimental results were consistent with most model predictions for browning: light penetration, benthic algal production, and zoobenthos biomass decreased, and pelagic nutrients and pelagic algal production increased with browning. Also consistent with expectations, warming had negative effects on benthic and pelagic algal biomass and weak effects on algal production and zoobenthos and zooplankton biomass. Inconsistent with expectations, browning had no effect on zooplankton and warming effects on fish depended on browning. The model is applicable also to nutrient‐rich systems, and we propose that it is a useful tool for the exploration of the consequences of different climate change scenarios for productivity and food web dynamics in shallow lakes, the worldwide most common lake type.     

Constraints on body size distributions: an experimental approach using a small-scale system

Holling’s (1992) proposition that discontinuities in biotic and abiotic processes generate structure in ecological systems is examined experimentally by imposing size-specific perturbations on marine sediment assemblages. Two kinds of perturbations were applied: organic enrichment and predation, each at two levels. Perturbations significantly affected the densities and relative abundance of the main invertebrate taxa and these effects were consistent with the known effects of enrichment and predation. However, there was little evidence of significant treatment effects on the overall benthic biomass or abundance size spectrum, supporting the contention that the spectrum is conservative and is probably constrained by habitat architecture. Received: 28 June 1999 / Accepted: 24 September 1999     

Upward cascading effects of nutrients: shifts in a benthic microalgal community and a negative herbivore response

We evaluated the effects of nutrient addition on interactions between the benthic microalgal community and a dominant herbivorous gastropod, Cerithidea californica (California horn snail), on tidal flats in Mugu Lagoon, southern California, USA. We crossed snail and nutrient (N and P) addition treatments in enclosures on two tidal flats varying from 71 to 92% sand content in a temporally replicated experiment (summer 2000, fall 2000, spring 2001). Diatom biomass increased slightly (~30%) in response to nutrient treatments but was not affected by snails. Blooms of cyanobacteria (up to 200%) and purple sulfur bacteria (up to 400%) occurred in response to nutrient enrichment, particularly in the sandier site, but only cyanobacterial biomass decreased in response to snail grazing. Snail mortality was 2–5 times higher in response to nutrient addition, especially in the sandier site, corresponding to a relative increase in cyanobacterial biomass. Nutrient-related snail mortality occurred only in the spring and summer, when the snails were most actively feeding on the microalgal community. Inactive snails in the fall showed no response to nutrient-induced cyanobacterial growths. This study demonstrated strongly negative upward cascading effects of nutrient enrichment through the food chain. The strength of this upward cascade was closely linked to sediment type and microalgal community composition.     

Influences of Multi-Scale Habitat on Metabolism in a Coastal Great Lakes Watershed

Spatial heterogeneity influences ecosystem structure and function across multiple habitat scales. Although primary production and respiration are fundamental to energy cycling in aquatic ecosystems, we know relatively little about how habitat scale influences metabolism. In this study, we adopted a multi-scale habitat approach to evaluate primary production and respiration in a coastal Great Lakes watershed that is experiencing pressure from past, present, and anticipated future human activities. We divided the watershed into five macrohabitats (stream, river, wetland, drowned-river mouth lake, and Great Lake), two mesohabitats (benthic and water column), and four microhabitats (periphytic substrates: sand/sediment, rock, wood, and plant) for evaluation of spatial patterns and synchrony in metabolism. Factors that influenced patterns of metabolism were scale dependent. Algal biomass strongly influenced spatial patterns in metabolism at the meso- and microhabitat scales; greater algal biomass translated to higher areal-specific and lower chlorophyll-specific metabolism at benthic mesohabitat and sand/sediment and rock microhabitats. Benthic metabolism overwhelmed water column metabolism, irrespective of location or time of year. Watershed position was important at the macrohabitat scale, with greater overall metabolism in macrohabitats located lower in the watershed. Average synchrony in metabolism rates was greatest at the macrohabitat scale, suggesting metabolic patterns that are evident at finer scales may become integrated at coarser scales. Our results (1) show that spatial and temporal patterns in metabolism are shaped by factors that are dependent upon habitat scale; (2) highlight the importance of benthic productivity across habitat and season; and (3) suggest that hydrologic connectivity strongly influences ecosystem processes, although physical factors can affect these responses as evidenced by the low levels of synchrony between Lake Michigan and the other macrohabitats.     

Ecological regime shifts in salinised wetland systems. II. Factors affecting the dominance of benthic microbial communities

This paper is the second in a pair investigating potential mechanisms for ecological regime change in salinising wetlands. The first paper in this series focused on the responses of the salt-tolerant submerged macrophyte community to salinity. In this second paper, we investigated some of the environmental conditions required for initiation and dominance of benthic microbial communities using a combination of experimental and observational data. Two experiments were carried out. One investigated the importance of prior establishment of benthic microbial communities on their ability to maintain prevalence over macrophyte colonisation (‘persistence’ experiment), while the other investigated hydrology and its effect on sediment perturbation, potential nutrient release and subsequent benthic microbial community establishment (‘flooding’ experiment). The ‘persistence’ experiment measured the biomass of benthic microbial communities and emergence of macrophytes from sediments kept either wet or dry for 4 weeks then flooded at a range of salinities. Benthic microbial biomass was similar across all of the salinities tested (15, 45 and 70 ppt), with a slight increase at higher salinities, suggesting that none of these limited benthic microbial community development. Pre-wetting of sediments usually increased benthic microbial community biomass and reduced macrophyte germination, but the latter was attributed to the presence of anoxic sediments rather than the increased benthic microbial community biomass. Germinating macrophytes emerged through benthic microbial communities or dense heterotrophic bacterial blooms, demonstrating that they could become dominant even when another community was already established. Field data supported these results, suggesting that the development of benthic microbial communities is not limited by salinity alone, but includes other factors, such as the water regime. In the ‘flooding’ experiment, the largest differences in nutrient concentrations ultimately lay between the pre-wet and pre-dry treatments (due to the greater release of nutrients and development of anoxia in the latter) rather than those subjected to fast versus slow flooding. In response to this, highest benthic microbial community biomass was in treatments with pre-wet sediment, corresponding with lower phytoplankton biomass.     

Environmental factors accounting for benthic macroinvertebrate assemblage structure at the sample scale in streams subjected to a gradient of cattle grazing

Macroinvertebrate assemblages were related to environmental factors that were quantified at the sample scale in streams subjected to a gradient of cattle grazing. Environmental factors and macroinvertebrates were concurrently collected so assemblage structure could be directly related to environmental factors and the relative importance of stressors associated with cattle grazing in structuring assemblages could be assessed. Based on multivariate and inferential statistics, measures of physical habitat (% fines and substrate homogeneity) had the strongest relationships with macroinvertebrate assemblage structure. Detrital food variables (coarse benthic and fine benthic organic matter) were also associated with assemblage structure, but the relationships were never as strong as those with physical habitat measures, while autochthonous food variables (chlorophyll a and epilithic biomass) appeared to have no association with assemblage structure. The amount of variation explained in taxa composition and macroinvertebrate metrics is within values reported from studies that have examined macroinvertebrate metric–sediment relationships. The % Coleoptera and % crawlers had consistent relationships with % fines during this study, which suggests they may be useful metrics when sediment is a suspected stressor to macroinvertebrate assemblages in Blue Ridge streams. Findings from this study also demonstrate the importance of quantitative sampling through time when research goals are to identify relationships between macroinvertebrates and environmental factors.     

Habitat structure mediates biodiversity effects on ecosystem properties

Much of what we know about the role of biodiversity in mediating ecosystem processes and function stems from manipulative experiments, which have largely been performed in isolated, homogeneous environments that do not incorporate habitat structure or allow natural community dynamics to develop. Here, we use a range of habitat configurations in a model marine benthic system to investigate the effects of species composition, resource heterogeneity and patch connectivity on ecosystem properties at both the patch (bioturbation intensity) and multi-patch (nutrient concentration) scale. We show that allowing fauna to move and preferentially select patches alters local species composition and density distributions, which has negative effects on ecosystem processes (bioturbation intensity) at the patch scale, but overall positive effects on ecosystem functioning (nutrient concentration) at the multi-patch scale. Our findings provide important evidence that community dynamics alter in response to localized resource heterogeneity and that these small-scale variations in habitat structure influence species contributions to ecosystem properties at larger scales. We conclude that habitat complexity forms an important buffer against disturbance and that contemporary estimates of the level of biodiversity required for maintaining future multi-functional systems may need to be revised.     

Size-related auto-fragment production and carbohydrate storage in auto-fragment of <Emphasis Type="Italic">Myriophyllum spicatum</Emphasis> L. in response to sediment nutrient and plant density

Size-related asexual reproduction of submersed macrophytes is still poorly understood. Here, we investigate how size-related auto-fragmentation in Myriophyllum spicatum L. responds to sediment nutrients and plant density. An experiment was carried out with sediments containing two different nutrient levels and with two levels of plant density. The results show that sediment nutrients and plant density brought about a strong dependency of auto-fragment production and the amount of total non-structural carbohydrate (TNC) storage in auto-fragments on individual plant size (total plant biomass). However, these two factors acted differently on size dependency. Sediment nutrients positively affected auto-fragment production and the amount of TNC in auto-fragments of M. spicatum. High concentrations of sediment nutrients significantly increased these two traits in absolute value and the value relative to plant size. Although the auto-fragment biomass and the amount of TNC in auto-fragments did not differ between density treatments when plant size was considered, the absolute values of these two traits were much larger in the low plant density treatment than in the high plant density treatment, which suggested an indirect negative effect of plant density on the auto-fragmentation of M. spicatum. In addition, higher percentages of large auto-fragments (>100 mg) were produced by plants that grew in nutrient poor sediment and low plant density environment than plants in nutrient rich sediment and high plant density environment. These results do not solely highlight a size-dependent effect, but also a size-independent effect of auto-fragment production and the amount of TNC in auto-fragments of M. spicatum. Furthermore, such size-independent effects can be explained by the significant biomass partitioning differences and the similar TNC-concentrations in auto-fragments under different environmental conditions.