Long-term recovery dynamics determined by the degree of the disturbance - Ten years tracking of aquatic macroinvertebrate recolonisation after an industrial disaster (Red Sludge Disaster, Hungary).

A ten-year-long examination of macroinvertebrate community recovery was conducted following a catastrophic spill of highly alkaline red sludge (pH >13) into lowland streams. Our primary objective was to compare recovery patterns after coarse- and fine-grain disturbances, focusing on two aspects: i) trend analysis to reveal long-term changes of six community parameters, and ii) variation analyses to assess parameter changes over time. We conducted statistical analysis on long-term data series of macroinvertebrates obtained from quantitative samples collected at four sections with varying degrees of disturbance along the impacted stream sections. We developed a comprehensive theoretical framework comprising a series of sequential phases: Ramp-up, Overshoot, and Oscillation Phases. i) A trend analysis revealed that disturbances show a gradual recovery pattern, while variance analyses showed an asymptotic convergence to an equilibrium. ii) Evaluating these trends across phases unveiled that the initial recovery phase exhibited a steep trajectory, lasting 4-9 months, irrespective of disturbance severity. Coarse-grain disturbances induced a remarkable Overshoot phenomenon across all community metrics. The more severe the disturbance, the greater the height and duration of the Overshoot. Our results suggest that the presence or absence of Overshoot can serve as an indicator for coarse-grain disturbances in the context of large and infrequent disturbances (LID). The entire recovery process lasts for 2.5-3 years irrespective of the severity of the LID. In conclusion, a minimum survey duration of two and half years is deemed imperative to capture the phases of recovery, and changes associated with LID are not expected to extend beyond the three-year threshold. The theoretical framework, including Overshoot parameters, may assist future studies in comparing recovery patterns of different LID types. Furthermore, our theoretical framework is likely to be applicable to other groups of organisms given a sufficiently long monitoring of recovery, influenced also by the length of reproductive cycles.

Road crossings change functional diversity and trait composition of stream-dwelling macroinvertebrate assemblages.

Functional diversity is regarded as a key concept in understanding the link between ecosystem function and biodiversity, and is therefore widely investigated in relation to human-induced impacts. However, information on how the intersection of roads and streams (hereafter road crossings, representing a widespread habitat transformation in relation to human development), influences the functional diversity of stream-dwelling macroinvertebrates is still missing. The general aim of our study was to provide a comprehensible picture on the impacts of road crossing structures on multiple facets of the functional diversity of stream-dwelling macroinvertebrates. In addition, we also investigated changes in trait structure. Our research showed that road crossing structures had negative impacts on functional richness and dispersion; i.e., functional diversification. However, we found no significant impact on functional divergence and evenness components. We found a decrease in functional redundancy at road crossing structures. This indicates a reduced ability of the community to recover from disturbances. Finally, we found that road crossings drive stream habitat and hydrological changes in parallel with modification of the trait composition of stream-dwelling macroinvertebrate assemblages. All these results suggest that road crossings cause notable changes in the functional diversity of stream-dwelling macroinvertebrate assemblages.

Components of functional diversity revisited: A new classification and its theoretical and practical implications.

Functional diversity is regarded as a key concept for understanding the link between ecosystem function and biodiversity. The different and ecologically well-defined aspects of the concept are reflected by the so-called functional components, for example, functional richness and divergence. Many authors proposed that components be distinguished according to the multivariate technique on which they rely, but more recent studies suggest that several multivariate techniques, providing different functional representations (such as dendrograms and ordinations) of the community can in fact express the same functional component. Here, we review the relevant literature and find that (1) general ecological acceptance of the field is hampered by ambiguous terminology and (2) our understanding of the role of multivariate techniques in defining components is unclear. To address these issues, we provide new definitions for the three basic functional diversity components namely functional richness, functional divergence and functional regularity. In addition, we present a classification of presence-/absence-based approaches suitable for quantifying these components. We focus exclusively on the binary case for its relative simplicity. We find illogical, as well as logical but unused combinations of components and representations; and reveal that components can be quantified almost independently from the functional representation of the community. Finally, theoretical and practical implications of the new classification are discussed.

The type of forest edge shapes snail assemblages at forest-pasture transitions.

Semi-natural, nutrient-poor calcareous grasslands are local biodiversity hotspots that are increasingly threatened by land use intensification, abandonment, or indirect effects from adjacent habitats. The habitat quality of these grasslands is often influenced by neighbouring forests or intensively managed agricultural land. For example, shrubs encroaching on grassland reduce the sensitive habitat, but at the same time represent a new habitat type (transition zone at gradual forest edge). We investigated the effects of gradual and abrupt forest edges on the species richness, abundance, species composition, functional diversity and number of species of conservation importance (red-listed species) of land snail assemblages at forest-pasture transitions in the Jura Mountains, Switzerland. Forest edge type influenced the snail assemblages in different ways. Transition zones at abrupt forest edges had a higher species richness and more snail individuals than transition zones at gradual forest edges. Transition zones also differed in land snail species composition. At gradual forest edges, the transition zones contained some openland snail species, while those at abrupt forest edges had a similar species composition to the forest interior. Functional diversity was significantly higher for snails in the forests and transition zones at both abrupt and gradual edges than in pastures. In contrast, pastures and transition zones at both abrupt and gradual edges had a significantly higher number of red-listed snail species. Based on our findings, we recommend the creation of gradual forest edges through regular forest management practices, rather than through shrub encroachment into pasture, which could reduce the size of the threatened habitat.

Characterising functional strategies and trait space of freshwater macroinvertebrates.

We examined the functional strategies and the trait space of 596 European taxa of freshwater macroinvertebrates characterized by 63 fuzzy coded traits belonging to 11 trait groups. Principal component analysis was used to reduce trait dimensionality, to explain ecological strategies, and to quantify the trait space occupied by taxa. Null models were used to compare observed occupancy with theoretical models, and randomization-based analyses were performed to test whether taxonomic relatedness, a proxy of phylogenetic signal, constrains the functional trait space of freshwater macroinvertebrates. We identified four major strategies along which functional traits of the taxa examined show trade-offs. In agreement with expectations and in contrast to existing evidence we found that life cycles and aquatic strategies are important in shaping functional structure of freshwater macroinvertebrates. Our results showed that the taxonomic groups examined fill remarkably different niches in the functional trait space. We found that the functional trait space of freshwater macroinvertebrates is reduced compared to the range of possibilities that would exist if traits varied independently. The observed decrease was between 23.44 and 44.61% depending on the formulation of the null expectations. We demonstrated also that taxonomic relatedness constrains the functional trait space of macroinvertebrates.

Evaluating the biological validity of European river typology systems with least disturbed benthic macroinvertebrate communities.

Humans have severely altered freshwater ecosystems globally, causing a loss of biodiversity. Regulatory frameworks, like the Water Framework Directive, have been developed to support actions that halt and reverse this loss. These frameworks use typology systems that summarize freshwater ecosystems into environmentally delineated types. Within types, ecosystems that are minimally impacted by human activities, i.e., in reference conditions, are expected to be similar concerning physical, chemical, and biological characteristics. This assumption is critical when water quality assessments rely on comparisons to type-specific reference conditions. Lyche Solheim et al. (2019) developed a pan-European river typology system, the Broad River Types, that unifies the national Water Framework Directive typology systems and is gaining traction within the research community. However, it is unknown how similar biological communities are within these individual Broad River Types. We used analysis of similarities and classification strength analysis to examine if the Broad River Types delineate distinct macroinvertebrate communities across Europe and whether they outperform two ecoregional approaches: the European Biogeographical Regions and Illies' Freshwater Ecoregions. We determined indicator and typical taxa for the types of all three typology systems and evaluated their distinctiveness. All three typology systems captured more variation in macroinvertebrate communities than random combinations of sites. The results were similar among typology systems, but the Broad River Types always performed worse than either the Biogeographic Regions or Illies' Freshwater Ecoregions. Despite reaching statistical significance, the statistics of analysis of similarity and classification strength were low in all tests indicating substantial overlap among the macroinvertebrate communities of different types. We conclude that the Broad River Types do not represent an improvement upon existing freshwater typologies when used to delineate macroinvertebrate communities and we propose future avenues for advancement: regionally constrained types, better recognition of intermittent rivers, and consideration of biotic communities.

Resilience of spider communities affected by a range of silvicultural treatments in a temperate deciduous forest stand.

To secure the ecosystem services forests provide, it is important to understand how different management practices impact various components of these ecosystems. We aimed to uncover how silvicultural treatments affected the ground-dwelling spider communities during the first five years of a forest ecological experiment. In an oak-hornbeam forest stand, five treatments, belonging to clear-cutting, shelterwood and continuous cover forestry systems, were implemented using randomised complete block design. Spiders were sampled by pitfall traps, and detailed vegetation, soil and microclimate data were collected throughout the experiment. In the treatment plots spider abundance and species richness increased marginally. Species composition changes were more pronounced and treatment specific, initially diverging from the control plots, but becoming more similar again by the fifth year. These changes were correlated mostly to treatment-related light intensity and humidity gradients. The patchy implementation of the treatments induced modest increase in both gamma and beta diversity of spiders in the stand. Overall, spiders gave a prompt and species specific response to treatments that was by the fifth year showing signs of relatively quick recovery to pre-treatment state. At the present fine scale of implementation the magnitude of changes was not different among forestry treatments, irrespective of their severity.

Multi-species occupancy modeling provides novel insights into amphibian metacommunity structure and wetland restoration.

A fundamental goal of community ecology is to understand species-habitat relationships and how they shape metacommunity structure. Recent advances in occupancy modeling enable habitat relationships to be assessed for both common and rare species within metacommunities using multi-species occupancy models (MSOM). These models account for imperfect species detection and offer considerable advantages over other analytical tools commonly used for community analyses under the elements of metacommunity structure (EMS) framework. Here, we demonstrate that MSOM can be used to infer habitat relationships and test metacommunity theory, using amphibians. Repeated frog surveys were undertaken at 55 wetland sites in southeastern Australia. We detected 11 frog species from three families (Limnodynastidae, Myobatrachidae, and Pelodryadidae). The rarest species was detected at only one site whereas the most common species was detected at 42 sites (naive occupancy rate 0.02-0.76). Two models were assessed representing two competing hypotheses; the best-supported model included the covariates distance to the nearest site (connectivity), wetland area, presence of the non-native eastern mosquitofish (Gambusia holbrooki), proportion cover of emergent vegetation, an interaction term between Gambusia and emergent vegetation cover, and the proportion canopy cover over a site. Hydroperiod played no detectable role in metacommunity structure. We found species-habitat relationships that fit with current metacommunity theory: occupancy increased with wetland area and connectivity. There was a strong negative relationship between occupancy and the presence of predatory Gambusia, and a positive interaction between Gambusia and emergent vegetation. The presence of canopy cover strongly increased occupancy for several tree frog species, highlighting the importance of terrestrial habitat for amphibian community structure. We demonstrated how responses by amphibians to environmental covariates at the species level can be linked to occupancy patterns at the metacommunity scale. Our results have clear management implications: wetland restoration projects for amphibians and likely other taxa should maximize wetland area and connectivity, establish partial canopy cover, and eradicate Gambusia or provide aquatic vegetation to mitigate the impact of this non-native fish. We strongly advocate the use of MSOM to elucidate the habitat drivers behind animal occupancy patterns and to derive unbiased occupancy estimates for monitoring programs.

Multiple stressor effects on alpha, beta and zeta diversity of riverine fish.

We examined the effects of regional scale land use and local scale environmental and biotic stressors on alpha, beta and zeta diversities of native fish communities in wadeable streams and non-wadeable rivers in the Danube basin, Hungary. Relationships among land use and local scale environmental and biotic stressors were weak both in streams and rivers, suggesting that these stressors act relatively independently. Alpha diversity decreased strongly with increasing local scale environmental stressor intensity in rivers. On the contrary, its response to stressors was more obscure in streams, where the best-fit statistical model indicated the importance of the interaction between land use, local scale environmental and biotic stressors, while the secondly ranked model highlighted the negative impact of local scale environmental stressors. Analysis of variance using distance matrices provided evidence that stressors alone and in interactions explained compositional differences of pairs of study sites (beta diversity). Considering the degree of overall degradation, both local (alpha) and among-site (beta and zeta) diversity indices responded to increasing stressor intensity, generally negatively. Riverine fish communities showed higher degrees of similarity (lower beta and higher zeta) than stream fish communities. They also showed increasing similarity (i.e. homogenization) with increasing overall stressor intensity, unlike stream fish communities, which showed no relationship with overall stressor intensity. Our results suggest that the relationships between land use and local scale environmental and biotic stressors can be complex and so do their effects on biodiversity. While stressor specific indices can provide information on the role of specific stressors in some cases, the examination of overall stressor effects is needed to assess realistically the effects of anthropogenic disturbances on native fish diversity. Diversity indices that quantify among-site changes in species composition, such as measures of beta and zeta diversity, can be fruitful for better understanding the role of multiple stressors in structuring ecological communities.

Aerobic anoxygenic phototrophs are highly abundant in hypertrophic and polyhumic waters.

Aerobic anoxygenic phototrophs (AAPs) are a group of photoheterotrophic bacteria common in natural waters. Here, AAP abundance and contribution to total bacterial abundance and biomass were investigated to test whether the trophic status of a lake or content of coloured dissolved organic matter (CDOM) play a role in determining AAP distribution and abundance in shallow inland lakes, with special focus on hypertrophic and polyhumic waters. Twenty-six different shallow lakes in Hungary were monitored. AAP abundance and biomass were determined by epifluorescence microscopy. The lakes exhibit a broad range of CDOM (2-7000 mg Pt L-1) and phytoplankton biomass (2-1200 µg L-1 chlorophyll a concentration). Very high AAP abundance (up to 3 × 107 cells mL-1) was observed in polyhumic and hypertrophic shallow lakes. AAP abundance was influenced by phytoplankton biomass and CDOM content, and these effects were interrelated. As determined, 40 µg L-1 chlorophyll a and 52 mg Pt L-1 CDOM are threshold levels above which these effects have a synergistic relationship. Hence, the observed high AAP abundance in some soda pans is a consequence of combined hypertrophy and high CDOM content. AAP contribution was influenced by total suspended solids (TSS) content: the success of AAP cells could be explained by high TSS levels, which might be explained by the decrease of their selective grazing control.

Breeding system, shell size and age at sexual maturity affect sperm length in stylommatophoran gastropods.

BACKGROUND: Sperm size and quality are key factors for fertilization success. There is increasing empirical evidence demonstrating that sperm form and function are influenced by selective pressures. Theoretical models predict that sperm competition could favour the evolution of longer sperm. In hermaphrodites, self-fertilizing species are expected to have shorter sperm than cross-fertilizing species, which use sperm stored from several mating partners for the fertilization of their eggs and thus are exposed to intense sperm competition. We tested this hypothesis by comparing original data on sperm length in 57 species of simultaneously hermaphroditic stylommatophoran gastropods from Europe and South America with respect to the species' breeding system. We used 28S rRNA nuclear and COI mitochondrial sequence data to construct a molecular phylogeny. Phylogenetic generalized linear models were applied to examine the potential influence of morphological and life-history characters. RESULTS: The best-fit model revealed that the breeding system and age at sexual maturity influence sperm length in gastropods. In general, species with predominant cross-fertilization had longer sperm than species with predominant self-fertilization or a mixed breeding system. Across species with shells (snails), sperm length also increased with shell size. CONCLUSIONS: Our study provides evidence that sperm length in stylommatophoran gastropods is influenced by the risk of sperm competition, as well as by age at sexual maturity and shell size. This finding extends present knowledge of sperm evolution to a group of so far poorly studied simultaneous hermaphrodites.

A comparative analysis reveals weak relationships between ecological factors and beta diversity of stream insect metacommunities at two spatial levels.

The hypotheses that beta diversity should increase with decreasing latitude and increase with spatial extent of a region have rarely been tested based on a comparative analysis of multiple datasets, and no such study has focused on stream insects. We first assessed how well variability in beta diversity of stream insect metacommunities is predicted by insect group, latitude, spatial extent, altitudinal range, and dataset properties across multiple drainage basins throughout the world. Second, we assessed the relative roles of environmental and spatial factors in driving variation in assemblage composition within each drainage basin. Our analyses were based on a dataset of 95 stream insect metacommunities from 31 drainage basins distributed around the world. We used dissimilarity-based indices to quantify beta diversity for each metacommunity and, subsequently, regressed beta diversity on insect group, latitude, spatial extent, altitudinal range, and dataset properties (e.g., number of sites and percentage of presences). Within each metacommunity, we used a combination of spatial eigenfunction analyses and partial redundancy analysis to partition variation in assemblage structure into environmental, shared, spatial, and unexplained fractions. We found that dataset properties were more important predictors of beta diversity than ecological and geographical factors across multiple drainage basins. In the within-basin analyses, environmental and spatial variables were generally poor predictors of variation in assemblage composition. Our results revealed deviation from general biodiversity patterns because beta diversity did not show the expected decreasing trend with latitude. Our results also call for reconsideration of just how predictable stream assemblages are along ecological gradients, with implications for environmental assessment and conservation decisions. Our findings may also be applicable to other dynamic systems where predictability is low.

Optimization of a chemical attractant for Epicometis (Tropinota) hirta Poda.

In field trapping tests in Hungary cinnamyl alcohol (3-phenyl-2-propen-l-ol) and transanethole [(1-methoxy-4-(1-propenyl)benzene)] attracted significantly more adult Epicometis (Tropinota) hirta (Coleoptera, Scarabaeidae, Cetoniinae) when presented together in the same bait compared to the single compounds Best attraction was recorded by a 1:1 mixture. Addition of other common floral scent compounds, ie. 3-methyl eugenol, 4-methoxy-cinnamaldehyde, anisylacetone, beta-ionone, cinnamyl acetate, cinnamic aldehyde, eugenol, indole, 2-phenylethanol or phenylacetaldehyde did not influence catches. The binary cinnamyl alcohol/trans-anethole bait described in this study is recommended for use in traps of E. hirta for agricultural purposes.