Tibor Erős

Characterizing connectivity relationships in freshwaters using patch-based graphs

Spatial graphs in landscape ecology and conservation have emerged recently as a powerful methodology to model patterns in the topology and connectivity of habitat patches (structural connectivity) and the movement of genes, individuals or populations among these patches (potential functional connectivity). Most spatial graph’s applications to date have been in the terrestrial realm, whereas the use of spatially explicit graph-based methods in the freshwater sciences has lagged far behind. Although at first patch-based spatial graphs were not considered suitable for representing the branching network of riverine landscapes, here we argue that the application of graphs can be a useful tool for quantifying habitat connectivity of freshwater ecosystems. In this review we provide an overview of the potential of patch-based spatial graphs in freshwater ecology and conservation, and present a conceptual framework for the topological analysis of stream networks (i.e., riverscape graphs) from a hierarchical patch-based context. By highlighting the potential application of graph theory in freshwater sciences we hope to illustrate the generality of spatial network analyses in landscape ecology and conservation.

A measure for assessing functional diversity in ecological communities

Functional diversity is regarded as a key in understanding the link between ecosystem function and biodiversity, but its measurement is rather problematic. The two widely used continuous measures are the dendrogram-based measure (DBM) and the functional attribute diversity (FAD). In contrast to DBM, FAD does not require the knowledge of the entire species pool before the analysis, and hence FAD is a more ideal tool for measuring functional diversity. However, the original form of FAD and its variants have several undesirable properties. Here, we suggest a modified FAD (denoted by MFAD), which—as illustrated by artificial and actual data sets—allows calculating functional diversity without violating the twinning and monotonicity criteria such that the number of species collected is compensated for. These requirements are met by replacing the original species by so-called functional species and then by dividing FAD by the number of functional units. Accordingly, MFAD measures the dispersion of species in the functional traits space so that MFAD values for different communities can directly be compared if the same set of functional traits is used. Finally, using data of two freshwater communities (caddisfly and riverine fish), we evaluate the change of species richness and functional diversity in relation to sampling effort (sample unit size). We found that functional diversity is a better and more reliable community descriptor than species richness in a sense that it converges to the maximum faster in the function of sampling effort.

Human effects on ecological connectivity in aquatic ecosystems: Integrating scientific approaches to support management and mitigation

Understanding the drivers and implications of anthropogenic disturbance of ecological connectivity is a key concern for the conservation of biodiversity and ecosystem processes. Here, we review human activities that affect the movements and dispersal of aquatic organisms, including damming of rivers, river regulation, habitat loss and alteration, human-assisted dispersal of organisms and climate change. Using a series of case studies, we show that the insight needed to understand the nature and implications of connectivity, and to underpin conservation and management, is best achieved via data synthesis from multiple analytical approaches. We identify four key knowledge requirements for progressing our understanding of the effects of anthropogenic impacts on ecological connectivity: autecology; population structure; movement characteristics; and environmental tolerance/phenotypic plasticity. Structuring empirical research around these four broad data requirements, and using this information to parameterise appropriate models and develop management approaches, will allow for mitigation of the effects of anthropogenic disturbance on ecological connectivity in aquatic ecosystems.

Forest-Stream Linkages: Effects of Terrestrial Invertebrate Input and Light on Diet and Growth of Brown Trout (Salmo trutta) in a Boreal Forest Stream

Subsidies of energy and material from the riparian zone have large impacts on recipient stream habitats. Human-induced changes, such as deforestation, may profoundly affect these pathways. However, the strength of individual factors on stream ecosystems is poorly understood since the factors involved often interact in complex ways. We isolated two of these factors, manipulating the flux of terrestrial input and the intensity of light in a 2×2 factorial design, where we followed the growth and diet of two size-classes of brown trout (Salmo trutta) and the development of periphyton, grazer macroinvertebrates, terrestrial invertebrate inputs, and drift in twelve 20 m long enclosed stream reaches in a five-month-long experiment in a boreal coniferous forest stream. We found that light intensity, which was artificially increased 2.5 times above ambient levels, had an effect on grazer density, but no detectable effect on chlorophyll a biomass. We also found a seasonal effect on the amount of drift and that the reduction of terrestrial prey input, accomplished by covering enclosures with transparent plastic, had a negative impact on the amount of terrestrial invertebrates in the drift. Further, trout growth was strongly seasonal and followed the same pattern as drift biomass, and the reduction of terrestrial prey input had a negative effect on trout growth. Diet analysis was consistent with growth differences, showing that trout in open enclosures consumed relatively more terrestrial prey in summer than trout living in covered enclosures. We also predicted ontogenetic differences in the diet and growth of old and young trout, where we expected old fish to be more affected by the terrestrial prey reduction, but we found little evidence of ontogenetic differences. Overall, our results showed that reduced terrestrial prey inputs, as would be expected from forest harvesting, shaped differences in the growth and diet of the top predator, brown trout.

A proposed unified terminology of species traits in stream ecology

Traits-based community analyses are receiving increasing attention. However, consistent interpretation of empirical results and ecological understanding in stream ecology are limited by ambiguous terminology. Furthermore, the measurement scales used to analyze trait data, especially ordinal-scale data, are often inappropriately applied. We identify and discuss these shortcomings and offer a solution for an operative and algebraically correct treatment of traits and a unified nomenclature that facilitates direct comparison among traits-based studies. A unified terminology allows for logical translation among existing, alternative trait nomenclatures and should facilitate communication of research findings among stream ecologists and more directly connect stream traits-based research with general ecology.

Quantifying temporal variability in the metacommunity structure of stream fishes: the influence of non-native species and environmental drivers

Most studies characterize metacommunities based on a single snapshot of the spatial structure, which may be inadequate for taxa with high migratory behavior (e.g., fish). Here, we applied elements of metacommunity structure to examine variations in the spatial distributions of stream fishes over time and to explore possible structuring mechanisms. Although the major environmental gradients influencing species distributions remained largely the same in time, the best-fit pattern of metacommunity structure varied according to sampling occasion and whether or not we included non-native species in the analyses. Quasi-Clementsian and Clementsian structures were the predominant best-fit structures, indicating the importance of species turnover among sites and the existence of more or less discrete community boundaries. The environmental gradient most correlated with metacommunity structure was defined by altitude, area of artificial ponds in the catchment, and dissolved oxygen content. Our results suggest that the best-fit metacommunity structure of the native species can change in time in this catchment due to seasonal changes in distribution patterns. However, the distribution of non-native species throughout the landscape homogenizes the temporal variability in metacommunity structure of native species. Further studies are necessary from other regions to examine best-fit metacommunity structures of stream fishes within relatively short environmental gradients.

Functional diversity: a review of methodology and current knowledge in freshwater macroinvertebrate research

Although several studies have examined the functional diversity of freshwater macroinvertebrates, the variety of methodologies combined with the absence of a synthetic review make our understanding of this field incomplete. Therefore, we reviewed the current methodology for assessing functional diversity in freshwater macroinvertebrate research. Our review showed that most papers quantified functional diversity using biological traits, among which feeding habits were the most common traits probably due to the assumed links between feeding and ecosystem functions. A large number of diversity measures have been applied for quantifying functional diversity of freshwater macroinvertebrate assemblages, among which Rao’s quadratic entropy looks like the most frequent. In most papers, functional diversity was positively related to taxon richness, and functional redundancy was a key concept in explaining this correlation. Most studies detected strong influence of the environmental factors as well as human impact on functional diversity. Finally, our review revealed that functional diversity research is biased towards European running waters and is hindered by yet insufficient information on the autecology of macroinvertebrates.

Within-lake distribution patterns of fish assemblages: the relative roles of spatial, temporal and random environmental factors in assessing fish assemblages using gillnets in a large and shallow temperate lake

In this study, the relative role of spatio-temporal factors and associated environmental variables (water transparency and temperature) were quantified in relation to gillnet samples of fishes in a large and shallow lake (Lake Balaton, Hungary). Most of the variance (56·1%) in the relative abundance data (%) was related to the vertical segregation of fishes. This gradient substantially affected the catch per unit effort (CPUE) by number of the dominant species, the surface-oriented bleak Alburnus alburnus and the benthic common bream Abramis brama. It also influenced total CPUE, mean fish mass and species richness and diversity. At the lake level, horizontal habitat heterogeneity (i.e. littoral v. offshore) accounted for only 8·3% of the total variance in relative abundance data, but was important in structuring the CPUE of the ruffe Gymnocephalus cernua and the pikeperch Sander lucioperca. The longitudinal environmental gradient (i.e. lake basin), year and season of sampling, water transparency and temperature had significant effects on relative abundance only at the habitat level, but were also important components of variability of CPUE in some species at the lake level. As sampling schemes need to consider the main gradients in fish assemblage distributions, the use of surface and pelagic gillnets should be more intensively incorporated in the study and monitoring of fish assemblages in shallow lakes and lake habitats.

Does functional redundancy of communities provide insurance against human disturbances? An analysis using regional-scale stream invertebrate data

Human-induced reductions in species richness might alter the quality of ecosystem services when the remaining species are not able to substitute the functions provided by extirpated species. We examined how human disturbances (nutrient enrichment, land use intensification, instream habitat degradation and the presence of alien species) influence the species richness of stream invertebrates. Stream invertebrates (425 native species) were collected by kick and sweep sampling technique at 274 stream sites covering the entire area of Hungary. We measured the species richness, functional richness (i.e. number of unique functional roles provided by community members) and functional redundancy (i.e. the functional insurance of the community) using information on the feeding habits of each species. To remove the effect of natural variability, we tested the effect of stressors on the residuals of models relating species richness, functional richness and functional redundancy with natural environmental gradients. Our results showed that species richness was negatively influenced by instream habitat degradation and nutrient enrichment. Independent of the way of quantifying functional richness and functional redundancy, we found that functional richness is more sensitive to human impact than functional redundancy of stream invertebrates. The finding that a reduction of species richness is associated with a loss of unique functional roles (functional richness) is important for conservation issues, because the number of unique functional roles is usually regarded as driver of ecosystem functioning.

Dietary variability in fishes: the roles of taxonomic, spatial, temporal and ontogenetic factors

In spite of the general use of diet data in ecological research, still very little is known about the relative roles of spatial, temporal and biotic (e.g. taxonomic identity, size, sex) factors in dietary variability of fishes. Here, we applied canonical correspondence analysis and variation partitioning to examine the roles of taxonomic, annual, seasonal, lake basin, habitat and ontogenetic (standard length, LS) factors in the dietary variation of fishes in large and shallow Lake Balaton, Hungary. The analyses were performed at the assemblage (15 fish species) and the individual species levels, and based on high (24 fine resource categories) and low resolution (nine broad resource categories) diet data. As hypothesised, most of the explained variation related to interspecific differences, while the roles of sampling year, season, lake area, habitat and LS proved to be unexpectedly low at the assemblage level. In addition, no regularity was found in how the relative roles of these factors change between fish species. The high ratio of the unexplained variation suggests that individual variations in foraging strategies and resource use of fishes and unascertained stochastic processes had a strong influence on dietary variability both at the assemblage and the individual species levels.