András Specziár

Age- and size-specific patterns of heavy metals in the organs of freshwater fish Abramis brama L. populating a low-contaminated site.

Concentrations of cadmium, copper, mercury, lead and zinc were determined by atomic absorption spectrophotometry in the muscle, gill and liver of bream Abramis brama L. to study the relationship between the heavy metal load of fish and their age and size, and the seasonal variation of pollutant loads. Fish were collected from the Western basin of Lake Balaton (Hungary) in October 1999 and May 2000. The average metal concentrations of different organs varied in the following ranges: Cd 0.42-2.10; Cu 1.77-56.2; Hg 0.01-0.19; Pb 0.44-3.24; Zn 10.9-82.5 microg g(-1) dry weight. The highest Cd, Cu, Pb and Zn concentrations were detected in the gill or liver of fish, whereas the highest Hg concentrations were measured in the muscle. In the liver of bream for cadmium, copper and mercury the Pearson correlation analysis revealed positive associations related to age and size (length, net weight), as well as for the mercury load of all three investigated organs. In the muscle and gill the copper, lead and zinc concentrations, similarly to the lead and zinc concentrations of the liver, the associations related to age and size were negative. The correlations between the heavy metal concentrations of organs and the individual condition factors of fish samples proved to have opposite trends compared to those related to the age and size of fish. The seasonal variations in the heavy metal load of bream could be attributed rather to the seasonal change in the condition factor of fish than to variations in the pollutant load of the site.

Feeding strategy and growth of cyprinids in the littoral zone of Lake Balaton

The growth, diet and feeding strategy of five phytophil or phytolithophil species of Cyprinidae from the littoral habitats of Lake Balaton were investigated by examining their scales and foregut contents. The relationships between the total anterior radii of scales and the standard lengths were represented best by a power function for white bream Blicca bjoerkna, and linear functions for common bream Abramis brama, roach Rutilus rutilus and wild goldfish Carassius auratus gibelio, respectively. The backcalculated mean lengths for the first age groups of common bream, white bream and roach did not differ statistically from those obtained by direct observation on 0 group fish in late November 1995. Compared to other waters, common bream grows slowly, wild goldfish and roach rapidly, while the growth rate of white bream can be considered of medium speed in Lake Balaton. Common bream showed a generalized feeding pattern, consuming mainly chironomid larvae, detritus and Corophium curvispinun. Roach showed a clear shift between specialization for Dreissena and algae. Despite the dense population of D. polymorpha in the lake, the significance of the herbivorous adaptation of roach has not yet been made clear. Wild goldfish consumed mainly detritus but, in the open water region, it shifted to zooplankton. White bream preyed chiefly on D. polymorpha, but showed a mixed feeding pattern and utilized most of the available food resources. Carp had the most specialized feeding strategy and preyed mainly on D. polymorpha. According to the discriminant analysis, the five cyprinids exhibited significant food resource partitioning.

Updating water quality targets for shallow Lake Balaton (Hungary), recovering from eutrophication

The paper presents an overview about recovery of shallow Lake Balaton from eutrophication by assessing quantitative and qualitative changes in phytoplankton, zooplankton, and chironomids as a function of load reduction. The aim was to update the present water quality targets. The proposed targeting scheme supplements the existing one with a range of lake-specific ecological criteria. We conclude that simple targets (desired phytoplankton biomass and permissible load) are the best choice during the initial stage of eutrophication management, but more complex schemes including ecological criteria are needed to trace recovery when re-organization of the ecosystem takes place.

Spatial distribution and short-term changes of benthic macrofauna in Lake Balaton (Hungary)

Spatial and short-term changes in the composition and density of the macrobenthic fauna were studied in Lake Balaton, a large shallow lake in Central Europe (Hungary). Spatial differences were examined along five transects and short-term changes at two stations of different trophic state. The macrobenthos consisted almost exclusively of Tubificidae and Chironomidae of the species Chironomus gr. plumosus, Procladius choreus, Tanypus punctipennis, Microchironomus tener and Cladotanytarsus sp. The Oligochaeta biomass showed a significant short-term decrease from 1995 to 1998, and their average biomass proved to be 0.86 and 0.79 g WFW (wet formalin weight) m-2 in 1996 and 1997, respectively. Chironomidae showed significant spatial and short-term differences following variations in primary production. Higher primary production resulted in higher biomass (up to 153.4 g WFW m-2 and 9785 ind. m-2 in 1995) of Chironomus-Procladius community, while lower primary production resulted in a very low biomass and abundance of the predatory Procladius-Tanypus-Microchironomus community (average: 3.16 g WFW m-2 and 1311 ind. m-2 in 1997). As compared to other lakes of similar climatic conditions, the total production of oligohaetes and chironomids proved to be very low in 1996 and 1997 (49.2 and 54.1 KJ m-2 yr-1, respectively). The 90% of the average chironomid production of Lake Balaton in 1996 and 1997 belonged to Ch. gr. plumosus (5.63 and 5.13 g m-2 yr-1), P. choreus (4.3 and 3.47 g m-2 yr-1) and T. punctipennis (0.27 and 2.21 g m-2 yr-1).

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.

Feeding and competition of five cyprinid fishes in different habitats of the Lake Balaton littoral zone, Hungary

Abstract Carp, common bream, white bream, wild goldfish and roach are the most common large‐sized cyprinids in the littoral zone of Lake Balaton. Their cumulative average contribution to the littoral fish fauna is between 65.0% and 86.2% by weight but the relative abundance of each species differs by habitat. The diet of four of these five cyprinids significantly differed within and between all the three habitats examined The highest food‐niche overlap was detected between carp and white bream along the stony shoreline and in the reed grass stands (Horns Index). However, high food‐niche overlaps were common in the open water areas, where all these species compete for similar food types. According to our results, the interspecific competition among cyprinids is expected to increase with the implementation of shoreline regulations and the destruction of the reed‐grass habitat.

Size-dependent prey selection in piscivorous pikeperch Sander lucioperca and Volga pikeperch Sander volgensis shaped by bimodal prey size distribution

Prey size and species selection of pikeperch Sander lucioperca and Volga pikeperch Sander volgensis were investigated in relation to predator size in the shallow Lake Balaton, Hungary. Although their gape sizes were similar, S. lucioperca shifted to piscivory earlier and consumed fewer, but larger, prey than S. volgensis. Prey species preference of the two piscivores also differed. A bimodal prey size distribution resulted in a reclining sigmoid curve for the life span predator size to prey size relationship with inflexion points between 266 and 284 mm predator standard length (L(S) ) in S. lucioperca. In S. volgensis, as well as in S. lucioperca L(S) ≤ 350 mm, prey size increased monotonically with predator L(S) , following a power trend for all prey size variables. Prey depth to predator L(S) relationship varied significantly with prey species and prey number in both piscivores, and prey depth tended to be smaller in predators consuming more than one prey. Both predator species characteristically selected less active, benthic prey fishes in spite of their spiny fin rays, and small- and mid-sized predators selected for small prey. Relatively large prey were also eaten, however, especially by the smallest and largest S. lucioperca.

The relative importance of spatial and environmental processes in distribution of benthic chironomid larvae within a large and shallow lake

Although chironomids are popular model organisms in ecological research and indicators of bioassessment, the relative role of dispersal and environmental filtering in their community assembly is still poorly known, especially at fine spatial scales. In this study, we applied a metacommunity framework and used various statistical tools to examine the relative role of spatial and local environmental factors in distribution of benthic chironomid taxa and their assemblages in large and shallow Lake Balaton, Hungary. Contrary to present predictions on the metacommunity organisation of aquatic insects with winged terrestrial adults, we found that dispersal limitation can considerably affect distribution of chironomids even at lake scale. However, we also revealed the predominant influence of environmental filtering, and strong taxa–environment relationships were observed especially along sediment type, sediment organic matter content and macrophyte coverage gradients. We account that identified reference conditions and assemblages along with specified optima and tolerances of the abundant taxa can contribute to our understanding of chironomid ecology and be utilised in shallow lake bioassessment. Further, we propose that predictive models of species–environment relationships should better take into account pure spatial structuring of local communities and species-specific variability of spatial processes and environmental control even at small spatial scales.

Species diversity and abundance of plant-dwelling chironomids across hierarchical habitat and seasonal scales in the oxbow lakes of River Tisza, Hungary

In temperate regions, plant-dwelling chironomids can be considered as cyclic colonizers that inhabit seasonally ephemeral submerged and floating-leaved macrophytes. In this study, patterns of abundance and species richness of plant-dwelling chironomids were investigated within and among stands of three macrophyte species in oxbow lakes along the River Tisza (Hungary). Chironomids colonized macrophytes rapidly and most species occupied the habitat before it had completely developed in June. Assemblage structure and abundance of particular species varied considerably between plant species, oxbow lakes and summer months in a characteristic succession from June to August indicating the presence of dynamic selection mechanisms. Abundance of most species, except Endochironomus tendens, total chironomid abundance, within-sample and total species diversity were highest on the submerged plant Ceratophyllum demersum, and most species occurred at the beginning of the colonization succession, in June. The contribution of oxbow lakes (20.3%) and the month sampled (20.3%) to total chironomid diversity was higher than would be expected by chance alone and the contribution of within (19.7%) and between samples (13.5%) to total chironomid diversity was lower than would be expected by chance alone. We conclude that regional biodiversity conservation action plans should include multiple habitats. Moreover, the significant seasonal species turnover proved the need for seasonal sampling to assess accurately the total diversity of chironomids in the system.

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.