Detritus and nutrient dynamics in the shore zone of lakes: a review

The importance of detritus varies greatly among shore zones of lakes, but in a large majority of these regions detrital pathways prevail. Aside from a great spatial and seasonal variability, macrophytes and bottom sediments appear to be dominant stores of nutrients in these habitats. Macrophytes hold a central position in nutrient cycling in the shore-littoral lake zones. They are the main source of autochthonous detritus as they prevail in the total biomass of littoral organisms, and they are only rarely available as direct food of consumers. Various processes and interactions determine the role of macrophytes in nutrients dynamics. These are: the intensity of nutrient uptake and translocation, release of nutrients by healthy plants and from decomposing plants, exchange of elements between macrophytes and their periphyton, as well as interception of seston by macrophyte stands. Particular plant species differ in their time of dying and susceptibility to decomposition. The changes in decomposing material (size structure of particles and nutrient content) mean that detritus in various stages of decomposition differs in its role in trophic dynamics of shore-littoral lake zones. Several types of shore regions as regards detritus sources and retention level are discussed.     

抚仙湖不同类型岸带沉水植物分布及水体氮磷特征

2005年6-7月,通过野外断面采样并结合水质分析,对抚仙湖不同类型岸带沉水植物分布及水体氮磷特征进行了调查.结果表明:抚仙湖岸带分为4个类型,岩石陡峭岸带、冲积平缓岸带、河口岸带以及湖湾岸带;沉水植物种类分布最多出现在河口型岸带和湖湾岸带,均为9种;最低出现在岩石陡峭岸带,为5种;沉水植物生物量最高出现在冲积平缓岸带,为8 300 g·m-2,最低出现在岩石陡峭岸带,为2416 g·m-2;沉水植物分布最深为A1岩石陡峭岸带,平均水深8.5 m,最深达到11.0 m;最浅为B2冲积平缓岸带,平均水深1.9 m,最深为6 m;岸带水体综合氮磷指标,以D2湖湾岸带最高,总氮、总磷分别达到5.34和0.145 mg·L-;最低为A2岩石陡峭岸带,总氮、总磷分别为0.87和0.015 mg·L-1抚仙湖沉水植物对水体氮磷的吸收固定总量约为总氮为5.28×104 kg,总磷为7 500 kg.     

The influence of macrophytes on the spatial distribution of littoral rotifers

1. The effect of macrophytes on the spatial distribution of littoral rotifers was examined in Lake Rotomanuka, New Zealand (37°55'S, 175°19'E). Total rotifer abundances and those of abundant species, were compared between three macrophyte species, Myriophyllum propinquum , Eleocharis sphacelata and Egeria densa , and spatially across a littoral transect in relation to these species.
2. The abundances of many species, for example Euchlanis dilatata, Lecane closterocerca and L. lunaris, differed significantly between macrophyte species. More planktonic forms, Ascomorpha saltans , Keratella cochlearis and Synchaeta oblonga, however, showed no significant preference for macrophyte species.
3. Differences in rotifer abundances were evident even when different species of macrophyte grew in close proximity to one another, indicating that variations in physical and chemical conditions, which occur in the littoral of Lake Rotomanuka, could be largely discounted for much of the variation between macrophyte species.
4. Variation in rotifers between macrophytes was probably the result of a number of factors, including differences in macrophyte morphology, macrophyte age, epiphytic algal growths and the differential effects of predation by invertebrates and fish between macrophytes.
5. Variability of rotifer abundances spatially across the ecotone was less marked than between macrophyte species. The species of macrophyte occurring, and therefore the community composition and distribution of macrophyte species in the littoral, appears to be a major influence in the spatial structuring of rotifer communities in the littoral region of lakes.     

Distribution of macroinvertebrates in relation to physical and biological variables in the littoral zone of nine New Zealand lakes

Macroinvertebrates play a key role in the littoral zone of lakes. Macroinvertebrate community composition is closely linked to habitat conditions. To date, there have been few attempts to relate macroinvertebrates to habitat factors in lakes. In this study, nine mainly oligotrophic lakes from throughout New Zealand were surveyed for macroinvertebrates. The lakes were selected to represent a range of suspended sediment loading and lake level regimes. Within each lake, several sites were selected to provide a range of exposure to wave action. A multiple regression approach was taken to relate macroinvertebrate community composition and habitat characteristics. The results of the analysis suggest that the littoral zone of the lakes we studied could be divided into four general habitats. The first is the wave wash zone characterised by coarse substrates and macroinvertebrate taxa usually associated with lotic environments, such as Ephemeroptera and Plecoptera. The second habitat is associated with macrophytes and is limited at the top by wave action and at depth by light attenuation. In this zone, the snail Potamopyrgus antipodarum is dominant, along with Trichoptera and Odonata. At the base of the macrophytes is the detrital habitat characterised by fine, organic rich sediments and dominated by chironomids, oligochaetes and Trichoptera. At depths below the macrophyte zone, fine sediments are found, and bivalves such as the freshwater mussel Hyridella menziesi are common. While macroinvertebrate abundance can be highly variable, some general predictions of community structure can be made based on a few key environmental factors. Abundance of snails Odonata and Trichoptera was positively related to macrophyte biomass. Some macroinvertebrate groups such as oligochaetes, chironomids, snails and bivalves were more common in fine substrates, while Ephemeroptera were characteristic of coarse substrates. Detrital biomass was important for most of the macroinvertebrate groups studied showing a positive relationship for oligochaetes and Trichoptera and a negative relationship for Ephemeroptera and Plecoptera.     

The distribution of microcrustacea in the littoral zone of a freshwater lake

The distribution of microcrustacea in the water column, sediments and on different macrophyte species was examined in the littoral zone of Jack Lake, Nova Scotia, Canada. Large numbers of microcrustacea occurred in association with macrophytes, suggesting that this habitat should receive greater attention in future studies of microcrustacean numbers, biomass, and production. The relative abundance of different microcrustacea varied considerably among sediments, macrophytes and water column samples. Although microcrustacean species composition differed among macrophyte groups, consistent differences in absolute numbers per gram could not be demonstrated. Epiphytic microcrustacean community structure also varied among depth strata in Jack Lake. Few epiphytic and benthic microcrustacea migrated into the water column on a diurnal basis.     

Invertebrate necrobionts in the littoral zone in freshwater lakes of Karelia

The complex of necrobionts in the littoral zone in lakes of Karelia of different trophy states is considered. Forty-seven invertebrate species belonging to six classes and four types have been recorded on carcasses. The composition of necrobionts depends on the mass of carcasses, skin structure, and type of waterbody. Data on the rate of utilization are presented. The succession of necrobionts during the process of decomposition is considered.     

Structure and dynamics of zooplankton communities in the littoral zone of some North Carolina lakes

Species composition and seasonal, dynamics of zooplankton in the littoral zone were studied in three piedmont North Carolina lakes for one year. Nygaard's compound index indicated oligotrophic conditions in Belews Lake and eutrophic conditions in Salem Lake and Lake 150. A total of 32 species of zooplankton were found in the samples. Significantly fewer species of truely littoral zooplankton were present in the oligotrophic lake regardless of season (P < 0.05). Eutrophication appears to favor increased diversity in zooplankton of the littoral zone. Habitat heterogeneity, provided by a well developed littoral zone containing aquatic macrophytes, may be the major factor contributing to the maintenance of more taxa of zooplankton in eutrophic systems.Distinct taxa of Cladocera and Copepoda are present in the littoral zone, exhibiting seasonal dynamics that are largely different from the limnetic fauna. Such dynamics may result from predation by characteristically limnetic species of cyclopoid copepods which seek prey in the littoral zone. The numerical and seasonal abundance of predators and prey are inversely related and the true littoral taxa, represented in the present study by the genera Alona, Chydorus, Pleuroxus, Sida, Simocephalus, and Eucyclops, contained no predaceous species. The seasonal dynamics of these genera are very similar even in widely separated geographical regions, indicating that the factor(s) responsible operate independently of climatic and chemical variables. Intrazooplankton predation appears to be a more plausable explanation than segregation along depth which, although consistent with data from studies of limnetic waters, cannot be related to populations of zooplankton in shallow littoral areas.     

Spatial patterns of litter decomposition in the littoral zone of boreal lakes

1. We studied the patterns of litter decomposition in lake littoral habitats and investigated whether decay rates, as an integrating proxy for environmental conditions in the sediment, would co‐vary with net carbon dioxide (CO₂) exchange and methane (CH₄) efflux. These gas fluxes are known to be sensitive to environmental conditions. Losses in the mass of cellulose, root, rhizome and moss litter were measured during 2 years in boreal littoral wetlands in Finland and compared with published data on concurrently measured gas fluxes. Four study sites covered a range of sediment types and hydrological conditions. 2. Decomposition was not linearly related to the duration of flooding but depended on sediment type. Readily decomposable litter fractions, such as cellulose and rhizome litter, lost mass at a faster rate in marshes with a longer period of flooding but wide water level fluctuations that hinder establishment of a Sphagnum cover, than in peat‐forming fens. In marshes, the mean first‐year mass losses were 83–99% and 19–62% for cellulose and rhizomes, respectively. In fens, the respective losses were 40–53% and 33%. In the first year, the loss in the mass of the more recalcitrant root litter did not differ between sites (mean 19–30%) and moss litter lost no mass. 3. The estimated first‐year carbon loss from belowground litter was about 0.1–0.3 times ecosystem respiration and roughly similar to net carbon gas (CO₂, CH₄) efflux, suggesting that vascular plants and recent plant residues contribute substantially to ecosystem release of carbon gases. On the other hand, at least 40% of the mass of the belowground litter remained on a littoral site after the first 2 years of decomposition. Slow decomposition may indicate the accumulation of organic‐rich sediments. The accumulated carbon could explain the excess CO₂ release found in most littoral sites. In continuously inundated sites decomposition rates were similar to those in periodically flooded sites, but ecosystem‐atmosphere CO₂ exchange fell to close to zero. This discrepancy implies that the released CO₂ is dissolved in water and may be exported into the pelagic zone of the lake.     

湿地草本植物枯落物分解的影响因素

草本植物是湿地的主要植被类型之一,其枯落物是湿地有机物质的重要组成部分。本文概述了影响湿地草本植物枯落物分解的主要因素及研究进展,主要包括枯落物自身质量、生物因素、环境因素和人类活动与全球变化等。认为,枯落物质量是本质要素,生物是分解的主导因素,环境等外部因素起到重要作用。讨论了该领域有待深入研究的方向,指出要深入湿地枯落物分解机理的研究,加强多尺度、大环境梯度和多种影响因素的综合研究,加强全球变化对湿地枯落物分解的影响研究,增强实验方法研究等。     

Decomposition dynamics of aquatic macrophytes in the lower Atchafalaya, a large floodplain river

Decomposition of aquatic macrophytes can considerably influence carbon cycling and energy flow in shallow freshwater aquatic ecosystems. The Atchafalaya River Basin (ARB) is a large floodplain river in southern Louisiana that experiences a seasonal floodpulse and is spatially composed of a mosaic of turbid riverine and stagnant backwater areas. During two seasons, winter and fall of 1995, we examined decomposition of four common aquatic macrophytes in the ARB: water hyacinth (Eichhornia crassipes), arrowhead (Sagittaria platyphylla), coontail (Ceratophyllum demersum) and hydrilla (Hydrilla verticillata). To determine decay rates, we used litter bags of two mesh sizes (5 mm and 0.25 mm) and analyzed data with a single exponential decay model. Analysis of decay rates established several trends for aquatic macrophyte decomposition in the ARB. First, macrophytes decayed faster in fall than winter due to the effect of increased temperature. Second, macroinvertebrates were the primary decomposers of macrophytes in riverine sites and microbes were the primary decomposers in backwater areas. These trends may have been related to decomposer-habitat interactions, with well-oxygenated riverine sites more hospitable to invertebrates and backwater areas more favorable to microbes because of high organic inputs and reduced flow. Decay rates for macrophytes, ranked from slowest to fastest, were E. crassipes<S. platyphylla<C. demersum<H. verticillata. Slower decomposition of E. crassipes was probably a result of microbial inhibition by the waxy-cutin outer layer and low nutritional value. The accelerated decomposition of C. demersum and H. verticillata was most likely a function of the large surface area of the highly dissected leaves. Macroinvertebrate numbers were twice as high in riverine sites compared to backwater sites. In the winter, amphipods Gammarus spp. and Hyallela azteca composed a large percentage of the total density on detritus. In the fall, Caenis sp. was prevalent in the backwater habitat and dipterans were abundant in the riverine site. We investigated the microbial component involved in the decomposition of E. crassipes and S. platyphylla and found that the highest microbial respiration rates occurred early in the winter at the backwater site. Bacterial density in the winter on E. crassipes and S. platyphylla averaged 1.4×10⁶ cm^-2 after two days and decreased to 2.0×10⁵ cm^-2 after 28 d. Our results emphasized the importance of the microbial community in the decomposition of macrophytes in the ARB, especially in backwater habitats and in the early stages of decay.     

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.     

Removal of settled sediments and periphyton from macrophytes by grazing invertebrates in the littoral zone of a large oligotrophic lake

• 1 The resistance and resilience of littoral zone communities to sedimentation will depend both on the extent to which sediment deposition affects productivity, and on interactions within the communities. A series of hypotheses were set up and tested to examine interactions and feedback mechanisms among deposited sediments, periphyton, macrophytes and grazers in a large oligotrophic lake subject to fluctuating sediment loadings.
• 2 Although sediments incorporated into periphyton reduced light availability to macrophytes, periphytic algae were generally the dominant light absorbing component under natural conditions. When grazers were absent, both sediments incorporated in the periphyton and periphytic algal densities increased, and both were then important in reducing light available to macrophytes.
• 3 Grazing rate and assimilation efficiency for the dominant grazer, the prosobranch gastropod Potamopyrgus antipodarum, increased with increasing sediment content under natural lake conditions to reach a maximum at 10 mg sediment cm^?2.
• 4 An increase in sediment incorporation into periphyton films resulted in an increased grazing rate and hence grooming of sediments from macrophytes.
• 5 Grazing invertebrates can play a major role in maintenance of littoral communities by continuously grooming macrophyte hosts of periphytic algae and settled sediments.

     

Climate-related differences in the dominance of submerged macrophytes in shallow lakes

It has been suggested that shallow lakes in warm climates have a higher probability of being turbid, rather than macrophyte dominated, compared with lakes in cooler climates, but little field evidence exists to evaluate this hypothesis. We analyzed data from 782 lake years in different climate zones in North America, South America, and Europe. We tested if systematic differences exist in the relationship between the abundance of submerged macrophytes and environmental factors such as lake depth and nutrient levels. In the pooled dataset the proportion of lakes with substantial submerged macrophyte coverage (> 30% of the lake area) decreased in a sigmoidal way with increasing total phosphorus (TP) concentration, falling most steeply between 0.05 and 0.2 mg L⁻¹. Substantial submerged macrophyte coverage was also rare in lakes with total nitrogen (TN) concentrations above 1–2 mg L⁻¹, except for lakes with very low TP concentrations where macrophytes remain abundant until higher TN concentrations. The deviance reduction of logistic regression models predicting macrophyte coverage from nutrients and water depth was generally low, and notably lowest in tropical and subtropical regions (Brazil, Uruguay, and Florida), suggesting that macrophyte coverage was strongly influenced by other factors. The maximum TP concentration allowing substantial submerged macrophyte coverage was clearly higher in cold regions with more frost days. This is in agreement with other studies which found a large influence of ice cover duration on shallow lakes' ecology through partial fish kills that may improve light conditions for submerged macrophytes by cascading effects on periphyton and phytoplankton. Our findings suggest that, in regions where climatic warming is projected to lead to fewer frost days, macrophyte cover will decrease unless the nutrient levels are lowered.     

Edibility of fresh and decomposing macrophytes to three species of freshwater invertebrate herbivores

The edibility of two endemic Australian macrophytes (Ottelia ovalifolia and Triglochin procera) to three species of herbivorous stream invertebrates (Triplectides ciuskus and Notalina fulva: (Trichoptera: Leptoceridae) and Potamopyrgus niger (Gastropods: Hydrobiidae)) was investigated. Each macrophyte species was offered to the test animals in 5 different conditions: fresh, boiled, and three conditions of decomposition. In 5 out of 6 feeding trials, consumption rates of animals were greatest on decomposed tissues. Boiling increased edibility of both plant species to Triplectides larvae and of Triglochin to Notalina larvae. Boiled tissue was consumed least by Potamopyrgus. Analysis of nitrogen, phosphorus, and calorific content of macrophyte tissue and some terrestrial plant materials known to be eaten by the animals suggested that nutritional enrichment of decomposed and boiled tissues was unlikely to explain their greater edibility. Instead, the loss of anti-gustatory compounds is implicated.     

Effects of spring flood and water level draw-down on methane dynamics in the littoral zone of boreal lakes

1. The annual dynamics of methane (CH₄) in a temporarily flooded meadow, mire bank, lacustrine sedge fen, temporarily and continuously inundated sedge ( Carex sp.) and reed ( Phragmites australis ) marshes were studied from June to November in the humic mesoeutrophic Lake Mekrijärvi and in eutrophicated parts of the mesotrophic Lake Heposelkä in the southern part of East Finland. The effects of water level and temperature on littoral CH₄ fluxes were determined. Vegetation zonation along the moisture gradient, and associated CH₄ fluxes, were evaluated.
2. The CH₄ flux increased along the moisture gradient from –0.2 to 14.2 mg CH₄ m^–2 h^–1, and was highest in the permanently inundated marshes. The duration of anoxia in the sediment caused differences in the CH₄ flux. Estimated emissions for the period 1 June – 30 September in continuously inundated sparse reed and sedge marshes, drying sedge marsh, and lacustrine sedge fen were 13, 11 and 6 g CH₄ m^–2, respectively.
3. In continuously inundated vegetation, the fluxes were highest in late July/early August. The seasonal CH₄ flux pattern suggested that the fluxes were regulated by the supply of organic matter during the course of the summer and the water level. In the temporarily flooded zone, the seasonal CH₄ flux dynamics was greatly affected by changes in the lake water level, the fluxes being highest during the spring flood in early June.     

Role of macrophytes in the variability of zooplankton community structure in the littoral zone of shallow lakes

Distribution, diurnal variability, aggregation of zooplankton in the littoral zone of lakes and effect of various macrophyte species on the structure of its community are considered. It is shown that the horizontal migrations of zooplankton, both direct and reverse ones, are caused mainly by the pressure of fish. The effect of predacious zooplankton is less important and is directed mainly at small-sized species. The intensity of horizontal migrations of zooplankton decreases with depth, while the effect of shore avoidance is observed for the large-sized zooplankton species and is caused not only by the pressure of fish but also by other factors, most likely abiotic. The problem of interaction between macrophytes and zooplankton cannot be reduced to the role of macrophytes as a refuge. Allelopathic properties of macrophytes, competitive relations between separate zooplankton species in macrophyte thickets, as well as the effect of predacious invertebrates associated with macrophytes on zooplankton remain unclear. The role of macrophytes as a factor causing changes in hydrodynamic processes in the littoral regions of lakes is also unknown.     

Drag and reconfiguration of freshwater macrophytes

SUMMARY 1. Submerged freshwater macrophytes face large hydrodynamic forces in flowing waters in streams and on wave‐swept lake shores and require morphological adaptations to reduce the drag and the physical damage. This experiment studied five species of freshwater macrophytes and strap‐formed plastic leaves to test the predictions that: (i) increasing flexibility leads to greater reconfiguration and lower drag coefficients, (ii) flexible plants experience a steeper decline of drag coefficients with increasing water velocity than unflexible plants and (iii) plants mounted vertically on a horizontal substratum bend over in fast flow attaining a shielded position of low drag. 2. The results confirmed all three predictions. In fast flow, plants mounted upright on a horizontal platform gradually approached a position aligned with the flow, depending on their flexibility. In the range 8–50 cm s^?1 the deflection followed an interspecific negative linear relationship between log (tangent Φ) and velocity, where Φ represents the shoot angle normal to the horizontal level. Above 50 cm s^?1, further deflection was reduced perhaps by a combination of the elasticity and packing of shoots and the increasing lift generated by fast flow. 3. Drag coefficients of plants ranged between 0.01 and 0.1, typical of moderately to very streamlined objects. Drag coefficients declined log‐log linearly at increasing velocity, following negative slopes between ?0.67 and ?1.24 (median: ?1.0) because of reconfiguration and formation of a shielding canopy. Drag coefficients declined much less (median: ?0.55) for plants floating freely in the streaming water and which were capable of changing their shape but unable to form a shielding canopy. Drag coefficients declined even less for relatively unflexible plastic leaves (?0.30 to ?0.40), and they remained constant for stiff, bluff objects. 4. The experiments suggest that flow resistance of flexible, submerged macrophytes in natural streams may increase in direct proportion to water velocity because they form a shielding submerged canopy, and high water stages at peak flow may result in greater proportions of the water passing unimpeded above the canopy. In contrast, stiff amphibious and emergent reed plants should experience an increase of flow resistance with at least the square of velocity as reconfiguration is small and former aerial plant surfaces come into contact with the streaming water at higher water stages. Field experiments to test these predictions are urgently needed.     

The responses of littoral invertebrates to eutrophication-linked changes in plant communities

The decay of submerged macrophytes in lakes of high trophic level drastically limits the extent of habitat available to littoral invertebrates. The loss can be partially compensated by growth of filamentous algae. Our results show that macroinvertebrates typically associated with submerged macrophytes as well as planktonic crustaceans and rotifers occurred within algal mats at high densities.Aggregations of filamentous algae are usually short-term, with frequent appearances and disappearances. The rate of colonization of algal mats by invertebrates is rapid. In locations with a high degree of water exchange, animals colonize both living and decomposing algal mats at a similar rate, but in sheltered habitats, decomposing filamentous algae are colonized by a smaller number of animals.Comparison was made between the occurrence of invertebrate macrofauna on Chara spp., Potamogeton perfoliatus, P. lucens and Myriophyllum spicatum in several lakes. Although these macrophytes differ visibly in morphology and phenology, the number and composition of macroinvertebrates during summer was associated more closely with trophic state of a lake than with plant species.     

Benthic communities of exposed littoral sand-flats in eighteen Dutch lakes

Benthic communities of the exposed sandy littoral zone in eighteen Dutch lakes were sampled using a small, hand-held core (surface area 0.8 cm²). A community dominated by chironomid larvae, chydorid cladocerans, nematodes and copepods was found to be characteristic for this habitat, especially in large and/or deep lakes. Bray-Curtis similarities between sites were generally high (>70%). A hierarchical cluster analysis distinguished two clusters at a 75% similarity level only, varying in copepod and chydorid densities. The chironomidsCladotanytarsus andStictochironomus reached high densities in many lakes, whileRhynchotalona falcata, Monospilus dispar, Disparalona rostrata, Alona spp. andChydorus spp. (mainlyChydorus piger) were the most abundant chydorid taxa.     

Invertebrates of the littoral zone of Lake Glubokoe

This paper briefly reviews systematic, production and biocenotic studies on invertebrates of the littoral zone of Lake Glubokoe. It presents new data for a number of years on the spatial distribution and population dynamics of species (mainly cladocerans) constituting a zoocenosis confined to the lower surface of floating leaves of Nuphar luteum.