Jari Syväranta

An Empirical Evaluation of the Utility of Convex Hull and Standard Ellipse Areas for Assessing Population Niche Widths from Stable Isotope Data

Stable isotope analyses are increasingly employed to characterise population niche widths. The convex hull area (TA) in a δ13C–δ15N biplot has been used as a measure of isotopic niche width, but concerns exist over its dependence on sample size and associated difficulties in among-population comparisons. Recently a more robust method was proposed for estimating and comparing isotopic niche widths using standard ellipse areas (SEA), but this approach has yet to be tested with empirical stable isotope data. The two methods measure different kind of isotopic niche areas, but both are now widely used to characterise isotopic niche widths of populations. We used simulated data and an extensive empirical dataset from two fish populations to test the influence of sample size on the observed isotopic niche widths (TA and SEA). We resampled the original datasets to generate 5000 new samples for different numbers of observations from 5 to 80 to examine the statistical distributions of niche area estimates for increasing sample size. Our results illustrate how increasing sample size increased the observed TA; even sample sizes much higher than n = 30 did not improve the precision for the TA method. SEA was less sensitive to sample size, but the natural variation in our empirical fish δ13C and δ15N data still resulted in considerable uncertainty around the mean estimates of niche width, reducing the precision particularly with sample sizes n<30. These results confirm that the TA method is less appropriate for estimating population isotopic niche areas using small samples, especially when considerable population level isotope variation is expected. The results also indicate a need for caution when using SEA as a measure of trophic niche widths for consumers, particularly with low sample sizes and when the distribution and range for population isotope values are not known.

Evaluating the utility of stable isotope analyses of archived freshwater sample materials

We evaluated the potential utility of stable isotope analysis of tissues commonly archived by aquatic biologists. Previous studies with chemically preserved samples have shown contradictory results, which present an obstacle for the use of archived sample materials. We tested the effects of ethanol and formalin preservation on zooplankton and of ethanol on benthic macroinvertebrate δ13C and δ15N values. We found that neither formalin nor ethanol had a significant effect on δ13C and δ15N values of preserved zooplankton. Nor did ethanol significantly affect δ13C or δ15N values of macroinvertebrates. However, ethanol preservation slightly, but significantly decreased C:N ratios of both zooplankton and macroinvertebrates, probably reflecting some extraction of lipids. Overall, the effects of preservatives on δ13C and δ15N values that we observed were minor. We also compared δ13C and δ15N values analysed from roach scales and perch operculum bones with those analysed from muscle tissue. Decalcification of scales and operculum bones only slightly improved our comparison to muscle tissue δ13C and δ15N values. Decalcified scales had slightly higher δ13C and lower δ15N values. Similarly, decalcified operculum bones showed slightly increased δ13C and decreased δ15N values to those for fish muscle. Our results confirm that scales and operculum bones can provide a suitable proxy for fish muscle in isotope studies with minor correction. We conclude that various archived sample materials can indeed be used with confidence for historical reconstructions of freshwater food webs by stable isotope analysis.

Effects of predation pressure and resource use on morphological divergence in omnivorous prey fish

BackgroundBody shape is one of the most variable traits of organisms and responds to a broad array of local selective forces. In freshwater fish, divergent body shapes within single species have been repeatedly observed along the littoral-pelagic axes of lakes, where the structural complexity of near shore habitats provides a more diverse set of resources compared to the open-water zones. It remains poorly understood whether similar resource-driven polymorphism occurs among lakes that vary in structural complexity and predation pressure, and whether this variation is heritable. Here, we analyzed body shape in four populations of omnivorous roach (Rutilus rutilus) inhabiting shallow lakes. We tested the relationship between body shape, gradients of resources, predation pressure, and, in a subset of two lakes, diet composition. We used genome scans of 331 polymorphic AFLP markers to test whether there was a heritable component to the observed morphological diversification.ResultsBody shape differed among lakes and was significantly correlated to differences in predation pressure. Roach from the lake with highest predation pressure were most divergent from the average body shape of all populations, characterized by a more streamlined body and caudally inserted dorsal fins; features that facilitate predator escape. Surprisingly, diet composition was not associated with morphology. AFLP analysis revealed weak genetic differentiation among lakes and no isolation by distance (IBD). Outlier analysis detected three loci under positive selection with differing frequencies in the four populations. General linear models did not support an association of lake-specific genotypes with morphological variation.ConclusionBody shape was divergent among lakes, suggesting that processes previously reported from within single lakes may also be operating at the scale of whole lakes. We found no evidence for body shape being heritable, although sample size was small in these natural populations. Rather than habitat structure and diet, we conclude that predation had a stronger effect on the prevalence of local morphotypes. A variable morphotype facilitating the efficient uptake of a variety of spatially and temporarily scattered resources seems to be favored in these small aquatic systems.

Top consumer abundance influences lake methane efflux

Lakes are important habitats for biogeochemical cycling of carbon. The organization and structure of aquatic communities influences the biogeochemical interactions between lakes and the atmosphere. Understanding how trophic structure regulates ecosystem functions and influences greenhouse gas efflux from lakes is critical to understanding global carbon cycling and climate change. With a whole-lake experiment in which a previously fishless lake was divided into two treatment basins where fish abundance was manipulated, we show how a trophic cascade from fish to microbes affects methane efflux to the atmosphere. Here, fish exert high grazing pressure and remove nearly all zooplankton. This reduction in zooplankton density increases the abundance of methanotrophic bacteria, which in turn reduce CH4 efflux rates by roughly 10 times. Given that globally there are millions of lakes emitting methane, an important greenhouse gas, our findings that aquatic trophic interactions significantly influence the biogeochemical cycle of methane has important implications.

Altered energy flow pathways in a lake ecosystem following manipulation of fish community structure

We used carbon and nitrogen stable isotope analyses to assess the relative contributions from pelagic and littoral energy sources to higher trophic levels in a lake ecosystem before and after a major food web perturbation. The food web structure of the lake was altered when the population sizes of the most abundant fish species (small perch, roach and bream) were reduced during an attempt to improve water quality by biomanipulation. Fish removal was followed by dense year classes of young fish, which subsequently increased the utilisation of pelagic resources. This was reflected as a decrease in relative energy contribution from littoral sources and also led to more distinct pelagic and littoral food chains after fish removal. Community metrics calculated from stable isotope data indicated increased trophic diversity and occupied niche area, and reduced trophic redundancy in the food web. However, only minor changes were observed in fish trophic positions, although roach and pike occupied slightly lower trophic positions after fish removal. Despite the Jyväsjärvi ecosystem becoming more dependent on pelagic energy after fish removals, the littoral energy contribution was still substantial, particularly to certain fish species. Hence, our results support recent arguments for the importance of benthic production in lake ecosystems. More generally, our results illustrate how large-scale perturbations of food web structure can alter energy flow patterns through an entire ecosystem.

Contrasting response of two shallow eutrophic cold temperate lakes to a partial winterkill of fish

Food-web effects of winterkill are difficult to predict as the enhanced mortality of planktivorous fish may be counterbalanced by an even higher mortality of piscivores. We hypothesised that a winterkill in a clear and a turbid shallow lake would equalise their fish community composition, but seasonal plankton successions would differ between lakes. After a partial winterkill, we observed a reduction of fish biomass by 16 and 43% in a clear-water and a turbid small temperate lake, respectively. Fish biomass and piscivore shares (5% of fish biomass) were similar in both lakes after this winterkill, but young-of-the-year (YOY) abundances were higher in the turbid lake. Top-down control by crustaceans was only partly responsible for low phytoplankton biomass at the end of May following the winterkill in both lakes. Summer phytoplankton biomass remained low in the clear-water lake despite high abundances of YOY fish (mainly roach). In contrast, the crustacean biomass of the turbid lake was reduced in summer by a high YOY abundance (sunbleak and roach), leading to a strong increase in phytoplankton biomass. The YOY abundance of fish in shallow eutrophic lakes may thus be more important for their summer phytoplankton development after winterkill than the relative abundance of piscivores.

Bacterial community response to changes in a tri‐trophic cascade during a whole‐lake fish manipulation

Microbial communities play a key role in biogeochemical processes by degrading organic material and recycling nutrients, but can also be important food sources for upper trophic levels. Trophic cascades might modify microbial communities either directly via grazing or indirectly by inducing changes.in other biotic or in abiotic factors (e.g., nutrients). We studied the effects of a tri-trophic cascade on microbial communities during a whole-lake manipulation in which European perch (Perca fluviatilis) were added to a naturally fishless lake divided experimentally into two basins. We measured environmental parameters (oxygen, temperature, and nutrients) and zooplankton biomass and studied the changes in the bacterial community using next generation sequencing of 16S rRNA genes and cell counting. Introduction of fish reduced the biomass of zooplankton, mainly Daphnia, which partly altered the bacterial community composition and affected the bacterial cell abundances. However, the microbial community composition was mainly governed by stratification patterns and associated vertical oxygen concentration. Slowly growing green sulfur bacteria (Chlorobium) dominated the anoxic water layers together with bacteria of the candidate division ODI. We conclude that alterations in trophic interactions can affect microbial abundance, but that abiotic factors seem to be more significant controls of microbial community composition in sheltered boreal lakes.

Isotopic variability in lake littoral organisms presents a challenge for food web studies

Most studies of lake ecosystems, including those that use stable isotope analysis (SIA), have traditionally concentrated on the pelagic system (Beaudoin et al. 2001, Vander Zanden & Vadeboncoeur 2002). Hence, temporal, spatial, and interspecies variability in isotopic signatures of pelagic food web components (especially zooplankton) have been the focus of recent studies (e.g., Matthews & Mazumder 2003, 2007). Recent SIA studies have also clearly indicated, however, that benthic production and the littoral areas of lakes may contribute more than previously thought to the whole lake ecosystem energy budget (Hecky & Hesslein 1995, Vander Zanden & Vadeboncoeur 2002, Vander Zanden et al. 2006). Studies of lake food webs based on SIA often assume single isotope values to represent baseline values for pelagic and littoral food webs. These values may then be used as “end members” in isotope mixing models, such as to assess trophic positions or relative contributions from different energy sources. But such single values ignore temporal, spatial or especially inter-specific variation in stable isotope values for these primary consumers, as shown for pelagic consumers by Matthews & Mazumder (2003). Such variability may have considerable impacts on the outputs from mixing models and hence on the inferences drawn about food web structure. Lake littoral habitats have received much less attention in this respect, despite being important feeding areas for many fish species and therefore contributing significantly to the observed stable isotope signatures at higher trophic levels. Littoral habitats are also inherently more heterogeneous than pelagic habitats, and primary production from phytoplankton, periphyton, and macrophytes, combined with allochthonous sources of energy, potentially result in complex and variable isotopic signatures in littoral primary consumers (Syvaranta et al. 2006). We present stable isotope data for littoral invertebrate consumer populations (benthic invertebrates and zooplankton) from Lake Jyvasjarvi in central Finland and show how variation in isotope signatures between different littoral invertebrate consumers can be reflected at higher trophic levels.

Periphyton support for littoral secondary production in a highly humic boreal lake

Steep stratification and poor light penetration in highly humic lakes typically restrict oxygenated littoral areas to narrow lake margins. However, in some instances, surrounding floating vegetation mats can sustain highly productive periphyton and more diverse invertebrate communities than pelagic areas. Little is known about how these littoral food webs function or the extent to which the pelagic and littoral food webs are coupled. We added 15N-labeled ammonium nitrate to the floating moss mat surrounding the littoral zone of Mekkojärvi, a small highly humic and fishless lake in southern Finland. Our goal was to increase the δ15N values of periphyton to investigate the diets of littoral invertebrates and possible pelagic–littoral coupling in the lake. We divided the lake in 2 with a plastic curtain and added European Perch (Perca fluviatilis) to 1 basin while the other remained fishless. δ15N of periphyton and most littoral invertebrates increased well above the natural abundance levels. δ15N of pelagic Daphnia generally did not increase, except for a sudden and transitory increase in the basin where fish were introduced. Only one perch of the 33 recaptured following their introduction showed clearly increased δ15N. The lowest δ13C values were found in pelagic invertebrates. Most littoral invertebrates had values closer to those of periphyton, which clearly contributed significantly to the diets of most littoral invertebrate groups and was an important basal resource in the littoral food web. Chironomids and ephemeropterans had surprisingly low δ13C values, which may reflect inclusion in their diets of highly 13C-depleted methane-oxidizing bacteria, which were known to contribute to the diets of Daphnia in the lake. Our results indicate that the pelagic and littoral habitats are not strongly coupled in the absence of fish but that zooplanktivorous fish may increase coupling by driving zooplankton into the littoral zone to seek refuge from predation.

Contrasting response of two shallow eutrophic lakes to a partial winterkill of fish

Food-web effects of winterkill are difficult to predict as the enhanced mortality of planktivorous fish may be counterbalanced by an even higher mortality of piscivores. We hypothesised that a winterkill in a clear and a turbid shallow lake would equalise their fish community composition, but seasonal plankton successions would differ between lakes. After a partial winterkill, we observed a reduction of fish biomass by 16 and 43% in a clear-water and a turbid small temperate lake, respectively. Fish biomass and piscivore shares (5% of fish biomass) were similar in both lakes after this winterkill, but young-of-the-year (YOY) abundances were higher in the turbid lake. Top-down control by crustaceans was only partly responsible for low phytoplankton biomass at the end of May following the winterkill in both lakes. Summer phytoplankton biomass remained low in the clear-water lake despite high abundances of YOY fish (mainly roach). In contrast, the crustacean biomass of the turbid lake was reduced in summer by a high YOY abundance (sunbleak and roach), leading to a strong increase in phytoplankton biomass. The YOY abundance of fish in shallow eutrophic lakes may thus be more important for their summer phytoplankton development after winterkill than the relative abundance of piscivores.