Stefano Larsen

Anthropogenic modification disrupts species co‐occurrence in stream invertebrates

The question of whether species co-occurrence is random or deterministic has received considerable attention, but little is known about how anthropogenic disturbance mediates the outcomes. By combining experiments, field surveys and analysis against null models, we tested the hypothesis that anthropogenic habitat modification disrupts species co-occurrence in stream invertebrates across spatial scales. Whereas communities in unmodified conditions were structured deterministically with significant species segregation, catchment-scale conversion to agriculture and sediment deposition at the patch- or micro-habitat scale apparently randomized species co-occurrences. This shift from non-random to random was mostly independent of species richness, abundance and spatial scale. Data on community-wide life-history traits (body size, dispersal ability and predatory habits) and beta-diversity indicated that anthropogenic modification disrupted community assembly by affecting biotic interactions and, to a lesser extent, altering habitat heterogeneity. These data illustrate that the balance between predictable and stochastic patterns in communities can reflect anthropogenic modifications that not only transcend scales but also change the relative forces that determine species coexistence. Research into the effects of habitat modification as a key to understanding global change should extend beyond species richness and composition to include species co-occurrence, species interactions and any functional consequences.

Biodiversity change is uncoupled from species richness trends: consequences for conservation and monitoring

Global concern about human impact on biological diversity has triggered an intense research agenda on drivers and consequences of biodiversity change in parallel with international policy seeking to conserve biodiversity and associated ecosystem functions. Quantifying the trends in biodiversity is far from trivial, however, as recently documented by meta-analyses, which report little if any net change in local species richness through time. Here, we summarise several limitations of species richness as a metric of biodiversity change and show that the expectation of directional species richness trends under changing conditions is invalid. Instead, we illustrate how a set of species turnover indices provide more information content regarding temporal trends in biodiversity, as they reflect how dominance and identity shift in communities over time. We apply these metrics to three monitoring datasets representing different ecosystem types. In all datasets, nearly complete species turnover occurred, but this was disconnected from any species richness trends. Instead, turnover was strongly influenced by changes in species presence (identities) and dominance (abundances). We further show that these metrics can detect phases of strong compositional shifts in monitoring data and thus identify a different aspect of biodiversity change decoupled from species richness. Synthesis and applications: Temporal trends in species richness are insufficient to capture key changes in biodiversity in changing environments. In fact, reductions in environmental quality can lead to transient increases in species richness if immigration or extinction has different temporal dynamics. Thus, biodiversity monitoring programmes need to go beyond analyses of trends in richness in favour of more meaningful assessments of biodiversity change.

Weak Concordance between Fish and Macroinvertebrates in Mediterranean Streams

Although anthropogenic degradation of riverine systems stimulated a multi-taxon bioassessment of their ecological integrity in EU countries, specific responses of different taxonomic groups to human pressure are poorly investigated in Mediterranean rivers. Here, we assess if richness and composition of macroinvertebrate and fish assemblages show concordant variation along a gradient of anthropogenic pressure in 31 reaches across 13 wadeable streams in central Italy. Fish and invertebrate taxonomic richness was not correlated across sites. However, Mantel test showed that the two groups were significantly, albeit weakly, correlated even after statistically controlling for the effect of environmental variables and site proximity. Variance partitioning with partial Canonical Correspondence Analysis showed that the assemblages of the two groups were influenced by different set of environmental drivers: invertebrates were influenced by water organic content, channel and substratum features, while fish were related to stream temperature (mirroring elevation) and local land-use. Variance partitioning revealed the importance of biotic interactions between the two groups as a possible mechanisms determining concordance. Although significant, the congruence between the groups was weak, indicating that they should not be used as surrogate of each other for environmental assessments in these Mediterranean catchments. Indeed, both richness and patterns in nestedness (i.e. where depauperate locations host only a subset of taxa found in richer locations) appeared influenced by different environmental drivers suggesting that the observed concordance did not result from a co-loss of taxa along similar environmental gradients. As fish and macroinvertebrates appeared sensitive to different environmental factors, we argue that monitoring programmes should consider a multi-assemblage assessment, as also required by the Water Framework Directive.

Resource subsidies between stream and terrestrial ecosystems under global change.

Streams and adjacent terrestrial ecosystems are characterized by permeable boundaries that are crossed by resource subsidies. Although the importance of these subsidies for riverine ecosystems is increasingly recognized, little is known about how they may be influenced by global environmental change. Drawing from available evidence, in this review we propose a conceptual framework to evaluate the effects of global change on the quality and spatiotemporal dynamics of stream-terrestrial subsidies. We illustrate how changes to hydrological and temperature regimes, atmospheric CO2 concentration, land use and the distribution of nonindigenous species can influence subsidy fluxes by affecting the biology and ecology of donor and recipient systems and the physical characteristics of stream-riparian boundaries. Climate-driven changes in the physiology and phenology of organisms with complex life cycles will influence their development time, body size and emergence patterns, with consequences for adjacent terrestrial consumers. Also, novel species interactions can modify subsidy dynamics via complex bottom-up and top-down effects. Given the seasonality and pulsed nature of subsidies, alterations of the temporal and spatial synchrony of resource availability to consumers across ecosystems are likely to result in ecological mismatches that can scale up from individual responses, to communities, to ecosystems. Similarly, altered hydrology, temperature, CO2 concentration and land use will modify the recruitment and quality of riparian vegetation, the timing of leaf abscission and the establishment of invasive riparian species. Along with morphological changes to stream-terrestrial boundaries, these will alter the use and fluxes of allochthonous subsidies associated with stream ecosystems. Future research should aim to understand how subsidy dynamics will be affected by key drivers of global change, including agricultural intensification, increasing water use and biotic homogenization. Our conceptual framework based on the match-mismatch between donor and recipient organisms may facilitate understanding of the multiple effects of global change and aid in the development of future research questions.

Artificial Light at Night Affects Organism Flux across Ecosystem Boundaries and Drives Community Structure in the Recipient Ecosystem

Artificial light at night (ALAN) is a widespread alteration of the natural environment that can affect the functioning of ecosystems. ALAN can change the movement patterns of freshwater animals that move into the adjacent riparian and terrestrial ecosystems, but the implications for local riparian consumers that rely on these subsidies are still unexplored. We conducted a two-year field experiment to quantify changes of freshwater-terrestrial linkages by installing streetlights in a previously light-naive riparian area adjacent to an agricultural drainage ditch. We compared the abundance and community composition of emerging aquatic insects, flying insects, and ground-dwelling arthropods with an unlit control site. Comparisons were made within and between years using generalized least squares and a BACI design (Before-After Control-Impact). Aquatic insect emergence, the proportion of flying insects that were aquatic in origin, and the total abundance of flying insects all increased in the ALAN-illuminated area. The abundance of several night-active ground-dwelling predators (Pachygnatha clercki, Trochosa sp., Opiliones) increased under ALAN and their activity was extended into the day. Conversely, the abundance of nocturnal ground beetles (Carabidae) decreased under ALAN. The changes in composition of riparian predator and scavenger communities suggest that the increase in aquatic-to-terrestrial subsidy flux may cascade through the riparian food web. The work is among the first studies to experimentally manipulate ALAN using a large-scale field experiment, and provides evidence that ALAN can affect processes that link adjacent ecosystems. Given the large number of streetlights that are installed along shorelines of freshwater bodies throughout the globe, the effects could be widespread and represent an underestimated source of impairment for both aquatic and riparian systems.

Scale dependent biodiversity patterns in Mediterranean river catchments: a multi taxa approach

Although the important contribution of β-diversity to regional (γ) diversity is increasingly recognised, our understanding of how the spatial scaling of β-diversity differs among taxonomic groups is still limited, especially in dynamic lotic ecosystems. In this study, we assessed the difference in the partitioning of diversity at nested spatial scales, from reach to catchment, among riparian birds, fish and benthic macroinvertebrates in Mediterranean river systems. Fish and macroinvertebrates showed similar scaling patterns, with β-diversity always larger than expected by a random distribution of individuals at all spatial scales (among reaches, rivers and catchments), and local (α) diversity always lower than expected. Conversely, β-diversity of riparian birds appeared larger than expected only at the largest scale (among catchments), while local diversity did not differ from random expectation. For birds, however, results partly depended on the weighting of abundant and rare species. Although the relative contribution of β to γ-diversity did not differ substantially among groups (with multiplicative β representing five to six distinct communities), its deviation from random expectations showed marked differences indicating that functionally different groups exhibit distinct spatial patterns. This study is among the first to investigate scaling patterns in β-diversity across taxonomic groups with different ecological requirements and dispersal ability, and provides a holistic picture of riverine biodiversity. From a conservation perspective, the results suggest that, in these river systems, flexible conservation strategies are required in order to protect multiple taxonomic groups.

The Hydra regeneration assay reveals ecological risks in running waters: a new proposal to detect environmental teratogenic threats

The regenerative ability of Hydra vulgaris was tested as potential biomarker for the development of a new eco-toxicological index. The test is based on the regeneration rate and the aberration frequency of the columna (body and adhesive foot) after separation from head and tentacles by a bistoury. Particularly, 45 columnae were submerged in the rearing solution (that is Hydra medium) to have control, and 285 in potential contaminated waters to have treatments, collected from 19 sites along 10 rivers in central Italy. ANCOVA and chi-square tests were used to compare values from each site to a laboratory control. Subsequently the values on regeneration rate and aberration frequency were inserted in a double entry matrix, where the match of the two entries in the matrix provides the score of the proposed Teratogenic Risk Index (TRI). Each score corresponded to one of the 5 teratogenic risk classes, to which a risk level was associated: from 1 (no risk) to 5 (very high risk). On the whole, 32% of the studied sites were classified as no teratogenic risk while the remaining showed a variable risk level from low to very high. This study proposed for the first time an early warning system to detect the presence of teratogens in running waters, providing a rapid and cost-effective evaluation method. Therefore, TRI may contribute to initiate adequate measures to manage riverine habitats, and to monitor the running water teratogenic status. Specifically, this index may provide the opportunity to identify the disturbance sources and then to drive the decisions, together with competent authorities, on the catchment and landscape management and on the possible use of waters for urban, agricultural, and industrial activities, since they may show significant effects on the human health.

Student‐centred experiments with stream invertebrates

Field experiments are at the core of ecology, yet rarely find their way into education, probably due to a combination of technical challenges and risks associated with fieldwork. This may compromise student education and perception of the subject, making interesting, readily manageable experiments extremely important. We describe how the daily cycle of nocturnal mass movements by river invertebrates (‘drift’) can meet these challenges. The observation of this daily cycle, and simple manipulations using light (to test hypotheses about the triggers for drift) and fine sand (simulating degradation of stream habitats), requires minimal specialist equipment or expertise, yet affords numerous practical benefits and captures student interest through the novel fieldwork. Invertebrate drift illustrates numerous biological principles through investigations adjusted to suit different educational levels.

Integrating community assembly and biodiversity to better understand ecosystem function: the Community Assembly and the Functioning of Ecosystems (CAFE) approach

The research of a generation of ecologists was catalysed by the recognition that the number and identity of species in communities influences the functioning of ecosystems. The relationship between biodiversity and ecosystem functioning (BEF) is most often examined by controlling species richness and randomising community composition. In natural systems, biodiversity changes are often part of a bigger community assembly dynamic. Therefore, focusing on community assembly and the functioning of ecosystems (CAFE), by integrating both species richness and composition through species gains, losses and changes in abundance, will better reveal how community changes affect ecosystem function. We synthesise the BEF and CAFE perspectives using an ecological application of the Price equation, which partitions the contributions of richness and composition to function. Using empirical examples, we show how the CAFE approach reveals important contributions of composition to function. These examples show how changes in species richness and composition driven by environmental perturbations can work in concert or antagonistically to influence ecosystem function. Considering how communities change in an integrative fashion, rather than focusing on one axis of community structure at a time, will improve our ability to anticipate and predict changes in ecosystem function.

WFD ecological status indicator shows poor correlation with flow parameters in a large Alpine catchment

Since the implementation of the Water Framework Directive, the ecological status of European running waters has been evaluated using a set of harmonised ecological indicators that should guide conservation and restoration actions. Among these, the restoration of the natural flow regime (ecological flows) is considered indispensable for the achievement of the good ecological status, and yet the sensitivity of the current biological indicators to hydrologic parameters remains understudied. The Italian Star_ICMi well represents other similar WFD indicators; it is a macroinvertebrate-based multimetric index officially adopted to assess the ecological status of running waters at the national level. Recent legislation has also included the Star_ICMi as one of the indicators used to assess and prescribe ecological flows in river reaches regulated by water abstraction. However, the relationship between river hydrology and the Star_ICMi index is so far virtually unknown. Using data from the Trentino - Alto Adige Alpine region, we first assessed the relationship between the Star_ICMi and synthetic descriptors of the physico-chemical (LIMeco) and morphological (MQI) status of respectively 280 and 184 river reaches. Then, we examined the relation between the Star_ICMi and a set of ecologically-relevant hydrologic parameters derived from discharge time-series measured at 21 hydrometric stations, representing both natural and regulated river reaches. Although the Star_ICMi showed significant and linear relationships with the physico-chemical character and, slightly, with the morphological quality of the reaches, its response to flow parameters appeared weak or non-existent when examined with linear models. Mixed quantile regressions allowed the identification of flow parameters that represented limiting factors for macroinvertebrate communities and the associated Star_ICMi scores. In particular, the index showed ‘negative floors’ where lower values were observed in reaches with large temporal variation in flow magnitude as well as frequent low and high flow events. The modelled quantiles also tracked the transition of the index from acceptable to unacceptable conditions. The results suggest that while the central tendency of the Star_ICMi index is not strongly influenced by river flow character, some key flow parameters represent limiting factors that allow the index to reach its lowest values, eventually ‘pushing’ the site towards unacceptable ecological conditions. The identification of limiting flow parameters can aid the setting of hydrologic thresholds over which ecological impairment is likely to occur. Overall, however, results imply caution is needed in using biological indicator like the Star_ICMi for the quantitative assessment and design of ecological flows.