Patch- and reach-scale dynamics of a macrophyte-invertebrate system in a New Zealand lowland stream

Abundant growths of macrophytes are a common feature of streams in open lowland areas of New Zealand during summer, but the values of these to aquatic biota are poorly understood. We studied the temporal dynamics of, and associations amongst, elements of a macrophyte-invertebrate system to provide an improved information base for lowland stream management. The biomass of macrophytes increased significantly over the four quarterly sampling occasions from 43.8 g m^-2 in June to 370.8 g m^-2 in March; biomass was dominated by Egeria densa on all dates, except in December when Potamogeton crispus was dominant. We did not detect strong associations between epiphyton biomass and invertebrate abundance in our study, but this may reflect the fact that we sampled loosely-adhering epiphyton on young, surface-reaching shoots whereas invertebrates were collected from macrophytes growing through the water column. Density of some invertebrate species per gram dry weight of plant material varied by macrophyte type, with the chironomids Tanytarsus vespertinus and Naonella forsythi displaying positive correlations with Egeria and Potamogeton biomass, respectively. The shrimp Paratya curvirostris accounted for 50% of phytophilous invertebrate biomass, with Chironomidae the only other group to comprise more than 9%. Abundance of total phytophilous invertebrates displayed a positive linear relationship with macrophyte biomass in a sample (0.1 m²), and a humped relationship with species richness, such that highest numbers of taxa occurred at macrophyte biomass levels around 400 g dw m^-2. Our study suggests that intermediate macrophyte biomass levels are likely to enhance macroinvertebrate biodiversity in sandy-bottomed lowland streams. This revised version was published online in July 2006 with corrections to the Cover Date.     

Unraveling assembly of stream biofilm communities

Microbial biofilms assemble from cells that attach to a surface, where they develop into matrix-enclosed communities. Mechanistic insights into community assembly are crucial to better understand the functioning of natural biofilms, which drive key ecosystem processes in numerous aquatic habitats. We studied the role of the suspended microbial community as the source of the biofilm community in three streams using terminal-restriction fragment length polymorphism and 454 pyrosequencing of the 16S ribosomal RNA (rRNA) and the 16S rRNA gene (as a measure for the active and the bulk community, respectively). Diversity was consistently lower in the biofilm communities than in the suspended stream water communities. We propose that the higher diversity in the suspended communities is supported by continuous inflow from various sources within the catchment. Community composition clearly differed between biofilms and suspended communities, whereas biofilm communities were similar in all three streams. This suggests that biofilm assembly did not simply reflect differences in the source communities, but that certain microbial groups from the source community proliferate in the biofilm. We compared the biofilm communities with random samples of the respective community suspended in the stream water. This analysis confirmed that stochastic dispersal from the source community was unlikely to shape the observed community composition of the biofilms, in support of species sorting as a major biofilm assembly mechanism. Bulk and active populations generated comparable patterns of community composition in the biofilms and the suspended communities, which suggests similar assembly controls on these populations.     

Macrophyte diversity and composition in relation to substratum characteristics in regulated and unregulated Danish streams

1. The objective of the present study was to examine how the physical stream environment in regulated and unregulated lowland streams affects the diversity and distribution of macrophyte communities. We analysed the abundance, distribution and composition of macrophytes, together with physical parameters, in seven regulated and seven unregulated unshaded Danish stream reaches. 2. Total macrophyte coverage was similar in the regulated and unregulated streams, but species richness and Shannon diversity were higher in the unregulated streams. Overall, we found fifty-two different species in the regulated stream reaches and sixty-two in the unregulated stream reaches. The spatial distribution of macrophytes on the stream bottom was more heterogeneous in the unregulated streams. 3. We found positive correlations between the coverage and diversity of macrophytes and the coverage of coarse-textured substratum types on the stream bottom, as well as between macrophyte coverage and diversity and substratum heterogeneity. We also found that the macrophytes were more heterogeneously distributed where substratum heterogeneity was greater. 4. The species growing both submerged and emergent were more abundant in the regulated streams, whereas species growing only submerged were more abundant in the unregulated streams. Species growing submerged, species growing both submerged and emergent, and species only growing emergent segregated differently in a canonical correspondence analysis ordination. The submerged species were primarily associated with coarser-textured substrata, whereas species growing both submerged and emergent, and species growing only emergent were associated with finer-textured substrata. 5. The most abundant species in the regulated streams, Sparganium emersum, accounting for almost one-third of the total macrophyte coverage, was primarily associated with clay and sandy bottom substrata, whereas the most abundant species in the unregulated streams, Batrachium peltatum, was primarily associated with gravel and stony substrata.     

Hyporheic community composition in a gravel-bed stream: influence of vertical hydrological exchange, sediment structure and physicochemistry

Summary 1. We studied the relative contributions of the magnitude and direction of vertical hydrological exchange, subsurface sediment composition and interstitial physicochemistry in determining the distribution of hyporheic invertebrates in the Kye Burn, a fourth order gravel‐bed stream in New Zealand. 2. In winter 2000 and summer 2001, we measured vertical hydrological gradient (VHG), dissolved oxygen, water temperature and water chemistry using mini‐piezometers, each installed in a different upwelling or downwelling zone. Next to every piezometer, a freeze core sample was taken to quantify the sediment, particulate organic matter and invertebrates. 3. Dissolved oxygen concentration at 25 cm was high on both occasions (>9 mg L^?1) but was higher in winter than summer. Interstitial water temperature was higher in down than upwellings and was substantially higher in summer than winter. Other features of the subsurface sediments and interstitial nitrate–nitrite concentrations were similar on both occasions and in up and downwellings. Interstitial ammonium and soluble reactive phosphorous concentrations were higher in winter than summer and ammonium was higher in up than downwelling areas. 4. The proportion of fine sediment (63 μm–1 mm), sediment heterogeneity and VHG accounted for the greatest proportion of variance in invertebrate distributions in both summer and winter. 5. The hyporheos was numerically dominated by early instar leptophlebiid mayfly nymphs and asellotan isopods. Water mites were a taxonomically diverse group with 13 genera. Taxonomic diversity (Shannon–Weaver), but not taxon richness, was higher in upwelling areas, reflecting lower numerical dominance by a few taxa in these locations. 6. Sediment composition (particularly the amount of fine sediments) and vertical hydrological exchange determined the composition and distribution of the hyporheos. Patchiness in these factors is important in planning sampling regimes or field manipulations in the hyporheic zone.     

Invertebrate colonization and processing of maple leaf litter in a forested and an agricultural reach of a stream

Processing of maple leaf (Acer saccharum Marsh) packs, their colonization by invertebrates and nutrient dynamics in leaves were investigated in a forested reach and agricultural reach of Canagagigue Creek, Ontario. Shredders, Pycnopsyche, and collectors, Ephemerella subvaria, Stenonema vicarium and Baetis were significantly more numerous in packs at the forest site than in packs at the agricultural site, whereas filter feeders, especially blackflies, were significantly more numerous in packs at the agricultural site. Weight loss of litter packs was nearly equivalent at the two sites. However, there were major differences in the mechanism of processing between the sites. Physical abrasion and microbial activity governed weight loss of maple leaf packs at the agricultural site, whereas processing was governed mainly by microbial and invertebrate activity and, to a much lesser extent, by physical abrasion at the forest site. Both shredders and collector species played an important role in the processing of leaf material at the forest site. Greater uptake of N and P (P<0.05 in spring) and higher C concentrations were observed in leaf packs at the forest site than the agricultural site. Therefore, the results support the concept of retention of nutrients in forested areas and their export in deforested (agricultural) areas. Findings also indicated that the processing of leaf litter is not an efficient means of monitoring changes in stream ecosystems since leaf processing is affected by many factors, particularly physical abrasion.     

Aquatic macrophytes in saline lakes of the Canadian prairies

Vascular macrophyte species richness decreases with increasing salinity. Only three species of submerged plants (Potamogeton pectinatus, Ruppia maritima, R. occidentalis) tolerate hypersaline waters (>50 g l^-1, total of ionic constituents). Eight emergent species occur in more saline habitats but only five (Scirpus maritimus var. paludosus, Distichlisstricta, Puccinellia nuttalliana, Scirpus americanus, Triglochin maritima) occur commonly over a range of saline lakes into the hypersaline category. Usually, species tolerant of high salinities are found over the entire saline spectrum and even extend into subsaline waters (<3 g l^-1) and thrive there. A major increase in the number of species occurs below 5 g l^-1. As the water recedes plants such as Salicornia rubra, Suaeda calceoliformes, Hordeum jubatum and Sonchus arvensis invade.Submerged angiosperm distribution is controlled by total ion concentration and substrate texture plays no apparent role. Although angiosperms normally grow in all kinds of substrates, they occupy coarse substrates in Wakaw lake because suitable fine substrates are densely colonized by charophytes. In this lake light limited growth occurs to a depth of 5% of surface light. Light was not limiting in Redberry Lake but angiosperm growth was limited to the upper 8 m (10% or more of surface light). Thermal stratification and depth (pressure) were probably limiting istead. In meromictic Waldsea Lake the depth of the chemocline (6 m, 5% surface light) delimits angiosperm growth.     

Connectivity and cladoceran species richness in a metacommunity of shallow lakes

SUMMARY 1. The shallow ponds of the nature reserve 'De Maten' form a metacommunity, in which individual ponds are highly interconnected via a system of overflows and rivulets. This study reports on the relations between cladoceran species richness and (a) connectivity patterns and (b) local environmental variables.
2. No relation was found between local species richness and three connectivity variables or dispersal pathways.
3. Spatial configuration was related to richness, but was confounded by environmental variables for 2 of 3 years. In those 2 years, there was a significant linear relation between Secchi disc depth and species richness, suggesting an important impact of the clearwater/turbid state alternative equilibria in shallow lakes in determining cladoceran richness. Only in the year in which environmental variables were unimportant did connectivity between the ponds influence species richness.
4. These results suggest that local environmental variables related to the clearwater/turbid state alternative equilibria in shallow lakes are important in determining cladoceran species richness. Connectivity and dispersal of individuals between the different ponds only act secondarily by increasing the general species richness within a pond through dispersal from ponds with different environmental conditions.     

Dynamic Properties of Recruitment,Expansion and Mortality of Macrophyte Patches in Streams

Streams are physically perturbed habitats with high demands on the dispersal and recruitment to maintain plant populations. Yet, little is known about these important demographic processes for stream plants. Therefore, we studied the monospecific vegetation of Callitriche cophocarpa in a small Danish lowland stream to determine: 1) the importance of drifting shoots and seeds for recruitment of plants, and 2) the influence of water flow, light availability and patch size on recruitment, growth and mortality processes. We found that the majority (about 90%) of new colonising patches of plant stands derived from drifting shoots being caught around protruding stones, while few developed from seeds. Many new patches were lost in the flowing water before roots became well established. Flow exposure of the patches resulted in the main growth taking place in the downstream direction. Combined areal cover of Callitriche patches on the stream bottom reached an upper limit of about 70%, probably because areal expansion above this threshold was constrained by strong shear forces and coarse substrata developing in the flow channels between the patches. We discuss why efficient shoot dispersal and vegetative growth documented here for Callitriche is an optimal plant strategy in flow-perturbed streams in contrast to the production of numerous small seeds but limited vegetative spread among ruderal plants in perturbed habitats on land.     

Spatial responses of hyporheic invertebrates to seasonal changes in environmental parameters

1. Spatial relationships between hyporheic invertebrates and subsurface water flow patterns, sediment characteristics, water physicochemical parameters and several possible food sources were compared over three seasons at one site beneath a riffle. Measures of food sources included particulate organic matter (POM), bacterial activity (aerobic respiration, nitrate respiration and mineralisation of organic nitrogen) and microbial abundance. 2. Patterns of water flow changed significantly over the 9‐month study period, from predominantly upwelling beneath the entire riffle in spring, to distinct differentiation between downwelling and upwelling zones in summer and autumn. Water physicochemical parameters changed accordingly, showing gradually weaker correlations with depth and stronger correlations with zone between spring and autumn. 3. Despite these changes, depth remained the strongest predictor of invertebrate richness, density and taxon composition throughout the study period. However, invertebrate distributions were secondarily correlated with water physicochemical parameters, and a minor gradient in invertebrate distributions between downwelling and upwelling zones became stronger from spring to summer. 4. The correlations between invertebrates and physicochemical parameters changed in both magnitude and direction with season. In spring, invertebrates showed a negative correlation with surface water infiltration, whereas in summer and autumn, the correlation was positive. Correlations were strongest in summer, when interstitial dissolved oxygen concentrations were lowest. 5. No relationships were found between hyporheic invertebrates and POM, microbial abundance or activity. This suggests that at this site, proximity to the streambed surface and physicochemical variables are more important than the abundance of food in controlling invertebrate distributions.     

Modelling nutrient cycles in relation to food web structure in a biomanipulated shallow lake

The modelPCLAKE describes the phosphorus and nitrogen cycles within a shallow lake ecosystem, including the sediment and a simplified biological food web. All components are modelled in a generalized way rather than a very detailed one. This model has been applied to Lake Zwemlust, a small biomanipulated lake in The Netherlands. Formerly, this highly eutrophic lake was dominated by cyanobacteria and devoid of macrophytes. Biomanipulation was carried out in 1987 by pumping-out of the water, removal of all fish, and refilling of the lake with seepage water. The lake was restocked with some rudd, pike, zooplankton and seedlings of macrophytes, and then monitored up to 1992. Macrophytes developed rather quickly and reached their maximum biomass during the six-years period in 1989. Despite the continuously high nutrient (N and P) loading, algal biomass remained low due to nitrogen limitation, caused by competition with the macrophytes. From 1990 onwards, the macrophytes declined again and a species shift occurred, following an increase of herbivorous birds on the lake and the development of herbivorous fishes.Model simulations grossly reproduced the observed developments in Lake Zwemlust before and after the biomanipulation measures. The existence of multiple steady states at the same trophic state and the possible shift between them could be simulated well. This study also demonstrates the interrelation between system structure and the distribution and cycling of nutrients. It is concluded, that within general boundary conditions set by the trophic state of the system, the food web structure determines the actual nutrient flows and the occurrence of nutrient limitations of the primary producers. It is shown that both aspects can be integrated in one mathematical model. The long-term stability of the macrophyte dominance in the lake is discussed.     

Nitrogen cycling and dynamics in a macrophyte‐rich stream as determined by a release

1. A tracer release study was conducted in a macrophyte‐rich stream, the River Lilleaa in Denmark. The objectives of the study were to compare uptake rates per unit area of by primary producers and consumers in macrophyte and non‐macrophyte habitats, estimate whole‐stream uptake rates of and compare this to other stream types, and identify the pathways and estimate the rate at which enters the food web in macrophyte and non‐macrophyte habitats. 2. Macrophyte habitats had four times higher primary uptake rates and an equal uptake rate by primary consumers per unit habitat area as compared to non‐macrophyte habitats. These rates represent the lower limit of potential macrophyte effects because the rates will be highly dependent on macrophyte bed height and mean bed height in the River Lilleaa was low compared to typical bed heights in many lowland streams. Epiphytes accounted for 30% of primary uptake in macrophyte habitats, illustrating a strong indirect effect of macrophytes as habitat for epiphytes. N flux per unit habitat area from primary uptake compartments to primary consumers was four times lower in macrophyte habitats compared to non‐macrophyte habitats, reflecting much greater biomass accrual in macrophyte habitats. Thus, we did not find higher N flux from macrophyte habitats to primary consumers compared to non‐macrophyte habitats. 3. Whole‐stream uptake rate was 447 mgN m^?2 day^?1. On a habitat‐weighted basis, fine benthic organic matter (FBOM) accounted for 72% of the whole‐stream uptake rate, and macrophytes and epiphytes accounted for 19 and 8%, respectively. 4. We had expected a priori relatively high whole‐stream N uptake in our study stream compared to other stream types mainly due to generally high biomass and the macrophyte’s role as habitat for autotrophic and heterotrophic organisms, but our results did not confirm this. In comparison with other release study streams, we conclude that nutrient concentration is the overall controlling factor for N uptake rates across streams, mostly as a result of high biomass of primary uptake compartments in streams with high nutrient concentrations in general and not in macrophyte streams in particular. 5. Our results indicate that macrophytes play an important role in the longer‐term retention of N and thus a decrease in net downstream transport during the growing season compared to streams without macrophytes, through direct and indirect effects on the stream reach. Direct effects are high uptake efficiency, low turnover rate (partly due to no direct feeding on macrophytes) and high longevity. An indirect effect is increased sedimentation of FBOM in macrophytes compared to non‐macrophyte habitats and streams which possibly also increase denitrification. Increased retention with macrophyte presence would decrease downstream transport during the growing season and thus the N loading on downstream ecosystems.     

Effects of logging on macroinvertebrate production in a sand-bottomed, low-gradient stream

1. Macroinvertebrate production and macrophyte growth were studied in logged and unlogged sections of a sand‐bottomed, low‐gradient, blackwater stream on the Coastal Plain of Virginia, U.S.A. A section of the catchment had been clear‐cut 3 years prior to sampling. No logging occurred in the upstream area of the catchment, which had experienced almost no land disturbance by humans for over 100 years. 2. A primary difference among the logged and unlogged sections of the stream was in the abundance of macrophytes. The combined biomass of Sparganium americanum and of Chara sp. was over 300‐times greater in the logged than the unlogged section. 3. Annual macroinvertebrate production in the sediment was higher in the unlogged section (41 g dry mass m^–2) than in the logged section (25 g m^–2). 4. Annual macroinvertebrate production on Sparganium was higher in the logged section (10 g m^–2 of plant surface area) than in the unlogged section (6 g m^–2). Annual production associated with Chara, which occurred only in the logged section, was 196 g m^–2 of stream bottom covered by this plant. 5. Whole‐stream annual macroinvertebrate production, calculated by summing habitat‐specific production that was weighted by habitat availability, was greater in the logged section (103 g m^–2) than in the unlogged section (41 g m^–2). Sediments supported 99% of the annual production in the unlogged section, whereas macrophytes supported 76% in the logged section. 6. Much of the additional macroinvertebrate production in the logged section was by collector‐filterers living on macrophytes. Production by collector‐gatherers was also greater in the logged section, whereas production by other functional feeding groups changed little with logging. 7. Although logging along high‐gradient, rocky streams also results in increased macroinvertebrate production, that increase often is stimulated by greater periphyton growth rather than the macrophyte growth observed in this low‐gradient stream.     

How do grazers affect periphyton heterogeneity in streams?

The effects of grazing by stream invertebrates on algal biomass and spatial heterogeneity were tested experimentally in flow-through microcosms with natural substrates (rocks). One experiment tested the effects of fixed densities of three species of grazers (the caddisfly Allomyia sp. and two mayflies, Epeorus deceptivus and Baetis bicaudatus) on periphyton. Baetis was tested with and without chemical cues from fish predators, which reduced grazer foraging activity to levels similar to the less mobile mayfly (Epeorus). Mean algal biomass (chlorophyll a; chl a) was reduced in grazer treatments compared to ungrazed controls, but there were no differences among grazer treatments. Algal heterogeneity (Morisita index) increased with grazer mobility, with the highest heterogeneity occurring in the Baetis-no fish treatment (most mobile grazer) and the lowest in the caddisfly treatment (most sedentary grazer). A second experiment used a three factorial design, and tested whether initial resource distribution (homogeneous vs. heterogeneous), Baetis density (high vs. low) and fish odor (present vs. absent) affected grazer impact on algal resources. Abundances of Baetis and chl a on individual rocks were recorded to explore the mechanisms responsible for the observed distributions of algae. Initial resource heterogeneity was maintained despite being subjected to grazing. Mean chl a was highest in controls, as in experiment I, and effects of Baetis on algal biomass increased with grazer density. There were no fish effects on algal biomass and no effects of grazer density or fish on algal heterogeneity. At the scale of individual rocks Baetis was unselective when food was homogeneously distributed, but chose high-food rocks when it was heterogeneously distributed. Results of these mechanistic experiments showed that Baetis can track resources at the scale of single rocks; and at moderate densities mobile grazers could potentially maintain periphyton distributions observed in natural streams.