Lars Peters

Stoichiometric mismatch between littoral invertebrates and their periphyton food

Abstract: Ecological stoichiometry is considered a key concept in understanding con-straints in energy transfer at the plant-herbivore interface. However, whether this con-cept is relevant for benthic freshwater ecosystems is not fully known. Therefore, afield survey was conducted in 2003 during the growing season in the littoral zone ofLake Constance, a large pre-alpine lake in central Europe. The aim was to assess tem-poral variation in the elemental stoichiometric composition in both herbivorous macro-invertebrates and their food resource, the periphyton in two different lakes. The peri-phyton showed large temporal and spatial variation in carbon, nitrogen, and phospho-rus content, with particularly high molar C:P ratios of up to 1225:1. Periphyton C:Pand C:N ratios were often high and constantly above the Redfield ratio that is consid-ered optimal for autotrophic growth. In contrast to the pronounced fluctuations in thenutrient ratios of their food resource, the herbivorous macroinvertebrates showed onlyvery little variation in their nutrient ratios, which indicated that they are homeostatic,i.e., physiologically restricted to a comparatively narrow range of C:P and C:N ratios.Distinct species-specific C : P and C : N ratios were found for different taxonomicgroups of macroinvertebrates, which indicated different requirements of optimal die-tary C:P and C:N ratios and which might influence the ability of the taxa to competefor limiting elemental nutrients. Considering the temporally very high C:P and C:Nratios of the periphytic resource and the very low ratios of the consumer body tissue,this stoichiometric mismatch is likely to constrain growth and reproduction of these lit-toral invertebrates. Therefore, the concept of stoichiometric food quality limitationmight also be applicable to the littoral food web in lakes.Key words: benthos, C:N:P ratio, ecological stoichiometry, herbivory, homeostasis,lake littoral, macroinvertebrates, mismatch, nutrient ratios, phosphorus.

Species distribution of free-living nematodes and other meiofauna in littoral periphyton communities of lakes

Recent studies on meiofaunal and nematode communities have focused on soft sediments in streams, lakes and marine environments. Despite a large number of studies dealing with periphyton, meiofaunal and nematode communities, on littoral hard substrates in lakes have not yet been investigated in detail. Therefore, epilithic communities with particular emphasis on nematode species composition, were analysed in 17 Swedish lakes differing greatly in size, depth, trophic status and epilithic biomass. Nematode abundance ranged from 2.3 to 161.5 cm −2 , and the abundance of nematodes relative to total meiofauna ranged from 20 to 77% (mean 53%). Fifty-eight nematode species were identified; species numbers varied from eight to 34 species per lake. The dominant species were Rhabdolaimus aquaticus , Punctodora ratzeburgensis , Eumonhystera dispar and Crocodorylaimus flavomaculatus . Deposit feeders dominated (71% of total fauna), followed by suction feeders (14%), epistrate feeders (12%) and chewers (3%). Of 3624 nematodes examined, 54% were juveniles, 35% females, 6% males and 5% gravid females. Multivariate analysis of the nematode species composition revealed significant differences in the community structures among lakes. This is the first study to show that meiofauna is a numerically abundant group within littoral periphyton communities in lakes, with nematodes representing the dominant group.

An efficient in situ method for sampling periphyton in lakes and streams

We present an efficient in situ sampling device that allows a simple and quantitative sampling of natural periphyton communities. The Brush Sampler is based on a previously developed sampler, which was improved by the addition of an external water supply, ball valve closing mechanism, and special exchangeable stiff brushes to solve previously reported problems of biomass overestimation and underestimation. The Brush Sampler was tested under ambient conditions in the field at four different sampling sites in a lentic and a lotic system and compared to the old sampler and control samples. The tests revealed a high inaccuracy of the old sampler and showed that biomass estimates (ash-free dry mass and chlorophyll-a content) obtained with the improved Brush Sampler did not significantly differ from biomass values determined from scrapings of control samples collected at the same sites. Therefore, our modified sampling device can be used as a tool for quantitative and qualitative epilithon community analyses.

Nematode Diversity in Terrestrial, Freshwater Aquatic and Marine Systems

This chapter is an attempt to bring together the studies of nematode diversity in terrestrial, freshwater aquatic and marine environments, and to synthesize these into some sort of general hypotheses. Such hypotheses can then be used as the bases for a range of other scientific activities, including biomonitoring and hypothesis testing. The effects of many of the variables in studies of nematode diversity, such as methods used, resolution or scope of studies, and measures of diversity are discussed first. With this background, studies on nematode diversity are then summarized and compared at a general level. Finally, hypotheses of the general patterns and processes of nematode diversity are discussed. These general patterns form the natural background for biomonitoring: to evaluate any effect of an environmental change one must know the situation expected without the change, including the expected magnitude and direction of the effects of environmental change.

Spatial variation of grazer effects on epilithic meiofauna and algae

Abstract Periphyton is a key component of shallow littoral zones of lakes and streams because it is an important source of primary production and a food resource for herbivores. Meiofauna are abundant in periphyton, but macroinvertebrate grazer (macrograzers) effects on periphytic meiofauna have not been studied so far. We used a spatially structured field experiment (hierarchical nested design consisting of 3 subsites at each of 3 sites) in Lake Erken (Sweden) to investigate the effect of macrograzers on epilithic meiofauna and algae in periphyton by controlling macrograzer access to littoral periphyton communities. Overall, we found a strong negative effect of macrograzer presence on algal biomass and some evidence for negative macrograzer effects on meiofaunal abundance and community composition. The impact of macrograzers on both algae and meiofauna were highly variable between sites and subsites. The largest spatial differences were for macrograzer effects on meiofaunal abundance and composition. We also investigated the ability of macrograzers to reduce spatial heterogeneity of periphyton biomass, but the presence of macrograzers did not alter the variation in algal biomass and associated meiofauna among replicates. We conclude that strong local variability in algal biomass and meiofauna abundance exists between neighboring sites even in the presence of strong overall macrograzer effects. This local variability could be based on factors known to cause spatial heterogeneity, such as hydrodynamics, nutrients, substrate characteristics (size, texture, exposure), or biotic interactions.

Epilithic communities in a lake littoral zone: the role of water-column transport and habitat development for dispersal and colonization of meiofauna

Abstract During the past 30 y, periphyton has been recognized as a key component in the benthos of lake littoral zones. However, the role of meiofauna-sized organisms living in the periphyton and mechanisms regulating and influencing those communities largely have been neglected, and the process of community development and colonization pathways of periphytic meiofauna are unknown in lakes. We studied these processes on littoral hard substrates in an oligotrophic lake. We manipulated the access of meiofauna to artificial hard substrates for a period of 57 d in a field experiment. In one treatment, the direct colonization pathway via active crawling was prohibited by elevating substrates into the water column. Development of the meiofaunal community was compared between elevated and nonelevated (control) treatments. In addition, relationships among the meiofaunal communities on the substrates in the field experiment and those in sediment traps and on natural hard substrates were examined. Periphyton biomass (chlorophyll a, total organic matter) and inorganic matter increased significantly in both experimental treatments throughout the experiment. Meiofauna colonized the substrates quickly and reached maximum densities of 107 ind./cm2. The initial colonization phase (2–8 d) was characterized by a great variation in meiofaunal community structure; rotifers and crustaceans were the most abundant groups. Communities became less variable during the experiment and resembled natural communities on hard substrates at the end of the experiment. In general, development of meiofaunal abundance and periphyton biomass was rapid and proceeded similarly in both treatments. Our results show that colonization of littoral hard substrates can be driven by water-column transport of meiofauna and point to the importance of this pathway for dispersal and colonization of hard substrates by meiofauna.

Community development of free-living aquatic nematodes in littoral periphyton communities

The development of an epilithic nematode community and the mode of colonisation was monitored over a 57-day period during spring in a field experiment in the littoral zone of a large oligotrophic lake. Two types of experimental units were used: one type prohibited direct colonisation via active crawling by elevating substrates into the water column and the second type had substrates placed on the lake bottom. Data from the two types of units and from nearby natural hard substrates were compared. The nematodes quickly colonised on the introduced substrates and reached maximum densities of 123 individuals per 10 cm2 after 57 days. Nematode densities on elevated and non-elevated substrates did not differ significantly. The nematode abundance and community structure showed a large initial variation, but became more stable over time and resembled the natural community structure at the end of the experiment. The maximum number of nematode species was reached after 2 weeks of colonisation, with a maximum species number on days 12 and 14 in the experimental units and on day 19 on the surrounding natural hard substrates. The five numerically dominant species, Eumonhystera vulgaris, Chromadorina bioculata, Eumonhystera filiformis, Chromadorina viridis and Daptonema dubium, accounted for most of the variation between the different communities. The results indicate that water-column transport was the main colonisation pathway of epilithic nematodes. The distribution of nematode species points to potential morphological adaptations of some species for persistence on hard substrates.

Grazing effects on nematodes and other components of the periphyton at three depths in Lake Erken

Abstract. The periphyton of stony hard substrates in the littoral zone of lakes is inhabited by an abundant and diverse algae and meiofauna assemblage that is regulated by grazing. The objective of our study was to examine the effect of macroinvertebrate grazers on periphytic algae and meiofaunal organisms at different depths in the littoral zone of Lake Erken (Sweden). To this end, a 2-factorial field experiment was carried out in which macrofaunal access to the periphyton was manipulated at 3 different water depths (30, 70, and 200 cm). Grazing effects on algal biomass varied among depths and were highest in both deeper zones. Compared to algal biomass, the abundance of meiofaunal organisms was less affected by grazing because macrograzers significantly reduced meiofauna only at the 200-cm depth. The organism mainly responsible for the depth-dependent differences in grazer effects was the snail Theodoxus fluviatilis, which dominated macrofaunal assemblage at depths where grazing effects were the strongest (70 and 200 cm). The abundance of T. fluviatilis was positively correlated with the proportions of reduced algal biomass and reduced meiofaunal abundance. Meiofaunal groups and individual nematode species responded differently to grazing, so our results further revealed that grazing macroinvertebrates affected the composition of the meiofaunal assemblage. However, meiofaunal and nematode assemblages differed more across depths than across grazing treatments, suggesting that other depth-related factors (e.g., hydrodynamics or light) also were important for structuring periphytic meiofaunal assemblages.