Jonathan D. Tonkin

Disentangling environmental drivers of benthic invertebrate assemblages: The role of spatial scale and riverscape heterogeneity in a multiple stressor environment

It is broadly acknowledged that freshwater ecosystems are affected by multiple stressors, but the relative importance of individual stressors in impairing riverine communities remains unclear. We investigated the impacts of multiple stressors, incorporating in-stream water quality, riparian and catchment land use and stream morphology, on riverine benthic invertebrate communities, while considering the spatial scales of factors and the heterogeneity of riverscapes. We performed a stepwise regression procedure linking 21 abiotic and 20 community metrics using Generalized Linear Models on data from 1018 river sites spread across Germany. High impact stressors (e.g., nutrients and water temperature) were identified for various community metrics. Both the combination of relevant stressors and their explanatory value differed significantly across streams of different sizes and ecoregions. In large rivers, the riparian land use was less important in determining community structure compared to lower order streams. Thus, possible mitigating effects of revegetated riparian buffer strips are likely to be overwhelmed by the influence of catchment-wide land use. Our results indicated substantial variability in stressors for the range of metrics studied, providing insight into potential target parameters for effective ecosystem management. To achieve long lasting successes in managing, protecting and restoring running waters, it is of vital importance to recognize the heterogeneity of riverscapes and to consider large-scale influences.

Seasonality and predictability shape temporal species diversity.

Temporal environmental fluctuations, such as seasonality, exert strong controls on biodiversity. While the effects of seasonality are well known, the predictability of fluctuations across years may influence seasonality in ways that are less well understood. The ability of a habitat to support unique, non-nested assemblages of species at different times of the year should depend on both seasonality (occurrence of events at specific periods of the year) and predictability (the reliability of event recurrence) of characteristic ecological conditions. Drawing on tools from wavelet analysis and information theory, we developed a framework for quantifying both seasonality and predictability of habitats, and applied this using global long-term rainfall data. Our analysis predicted that temporal beta diversity should be maximized in highly predictable and highly seasonal climates, and that low degrees of seasonality, predictability, or both would lower diversity in characteristic ways. Using stream invertebrate communities as a case study, we demonstrated that temporal species diversity, as exhibited by community turnover, was determined by a balance between temporal environmental variability (seasonality) and the reliability of this variability (predictability). Communities in highly seasonal mediterranean environments exhibited strong oscillations in community structure, with turnover from one unique community type to another across seasons, whereas communities in aseasonal New Zealand environments fluctuated randomly. Understanding the influence of seasonal and other temporal scales of environmental oscillations on diversity is not complete without a clear understanding of their predictability, and our framework provides tools for examining these trends at a variety of temporal scales, seasonal and beyond. Given the uncertainty of future climates, seasonality and predictability are critical considerations for both basic science and management of ecosystems (e.g., dam operations, bioassessment) spanning gradients of climatic variability.

Scale-dependent effects of river habitat quality on benthic invertebrate communities — Implications for stream restoration practice

Although most stream restoration projects succeed in improving hydromorphological habitat quality, the ecological quality of the stream communities often remains unaffected. We hypothesize that this is because stream communities are largely determined by environmental properties at a larger-than-local spatial scale. Using benthic invertebrate community data as well as hydromorphological habitat quality data from 1087 stream sites, we investigated the role of local- (i.e. 100 m reach) and regional-scale (i.e. 5 km ring centered on each reach) stream hydromorphological habitat quality (LQ and RQ, respectively) on benthic invertebrate communities. The analyses showed that RQ had a greater individual effect on communities than LQ, but the effects of RQ and LQ interacted. Where RQ was either good or poor, communities were exclusively determined by RQ. Only in areas of intermediate RQ, LQ determined communities. Metacommunity analysis helped to explain these findings. Species pools in poor RQ areas were most depauperated, resulting in insufficient propagule pressure for species establishment even at high LQ (e.g. restored) sites. Conversely, higher alpha diversity and an indication of lower beta dispersion signals at mass effects occurring in high RQ areas. That is, abundant neighboring populations may help to maintain populations even at sites with low LQ. The strongest segregation in species co-occurrence was detected at intermediate RQ levels, suggesting that communities are structured to the highest degree by a habitat/environmental gradient. From these results, we conclude that when restoring riverine habitats at the reach scale, restoration projects situated in intermediate RQ settings will likely be the most successful in enhancing the naturalness of local communities. With a careful choice of sites for reach-scale restoration in settings of intermediate RQ and a strategy that aims to expand areas of high RQ, the success of reach-scale restoration in promoting the ecological quality of communities can be greatly improved.

The next generation of site-based long-term ecological monitoring: Linking essential biodiversity variables and ecosystem integrity

Global change effects on biodiversity and human wellbeing call for improved long-term environmental data as a basis for science, policy and decision making, including increased interoperability, multifunctionality, and harmonization. Based on the example of two global initiatives, the International Long-Term Ecological Research (ILTER) network and the Group on Earth Observations Biodiversity Observation Network (GEO BON), we propose merging the frameworks behind these initiatives, namely ecosystem integrity and essential biodiversity variables, to serve as an improved guideline for future site-based long-term research and monitoring in terrestrial, freshwater and coastal ecosystems. We derive a list of specific recommendations of what and how to measure at a monitoring site and call for an integration of sites into co-located site networks across individual monitoring initiatives, and centered on ecosystems. This facilitates the generation of linked comprehensive ecosystem monitoring data, supports synergies in the use of costly infrastructures, fosters cross-initiative research and provides a template for collaboration beyond the ILTER and GEO BON communities.

Time is no healer: increasing restoration age does not lead to improved benthic invertebrate communities in restored river reaches.

Evidence for successful restoration of riverine communities is scarce, particularly for benthic invertebrates. Among the multitude of reasons discussed so far for the lack of observed effects is too short of a time span between implementation and monitoring. Yet, studies that explicitly focus on the importance of restoration age are rare. We present a comprehensive study based on 44 river restoration projects in Germany, focusing on standardized benthic invertebrate sampling. A broad gradient ranging from 1 to 25years in restoration age was available. In contrast to clear improvements in habitat heterogeneity, benthic community responses to restoration were inconsistent when compared to control sections. Taxon richness increased in response to restoration, but abundance, diversity and various assessment metrics did not respond clearly. Restoration age was a poor predictor of community composition and community change, as no significant linear responses could be detected using 34 metrics. Moreover, only 5 out of 34 tested metrics showed non-linear shifts at restoration ages of 2 to 3years. This might be interpreted as an indication of a post-restoration disturbance followed by a re-establishment of pre-restoration conditions. BIO-ENV analysis and fourth-corner modeling underlined the low importance of restoration age, but revealed high importance of catchment-scale characteristics (e.g., ecoregion, catchment size and land use) in controlling community composition and community change. Overall, a lack of time for community development did not appear to be the ultimate reason for impaired benthic invertebrate communities. Instead, catchment-scale characteristics override the effectiveness of restoration. To enhance the ecological success of future river restoration projects, we recommend improving water quality conditions and catchment-scale processes (e.g., connectivity and hydrodynamics) in addition to restoring local habitat structure.

Consistent effects of productivity and disturbance on diversity between landscapes

Productivity and disturbance have a strong role in determining diversity patterns in nature yet whether they operate individually or interact to determine diversity is unclear. Moreover, what effect land-use change has on this relationship has not been assessed. We tested whether land use influenced the relationship between productivity, disturbance and diversity, and assessed the fit of three productivity-disturbance-diversity models, using data from multiple samplings of 16 streams in two contrasting regions of the North Island of New Zealand. As the Dynamic Equilibrium Model (DEM) has received inconsistent support in all ecosystems and little favorable applications in lotic systems, we applied this along with two previously developed for stream communities. Although the community structure differed between the two regions, the response of taxonomic richness to productivity and disturbance was consistent. That is, richness was log-linearly related to productivity and declined monotonically with disturbance. However, there was no evidence of an interactive effect of productivity and disturbance. When accounting for density (rarefaction) the results were inconsistent, exhibiting no relationship with productivity but declining with disturbance. Our results suggest both the Death and Tonkin productivity-disturbance-diversity models are the most applicable in these communities where disturbance simply removes taxa and productivity controls the upper limit to richness.

Exploring stream communities in a tropical biodiversity hotspot: biodiversity, regional occupancy, niche characteristics and environmental correlates

Exploring and describing biodiversity and the mechanisms structuring it is fundamental to advancing ecology. This is particularly pertinent in understudied biogeographical regions, such as the Afrotropics, that are characterised by strong seasonal climatic shifts. We investigated the characteristics of stream biodiversity in the Niger Delta region of Nigeria, a tropical biodiversity hotspot, by examining patterns in 20 stream invertebrate communities across both the wet and dry seasons. For this, we took a multi-faceted approach accounting for the three levels of biodiversity (α, β and γ), including partitioning the nestedness and turnover components of β diversity, regional occupancy-abundance patterns, niche characteristics, and the environmental drivers of community structure. α diversity was low in these streams, with strong turnover between sites leading to high β diversity contributing to regional biodiversity, but there was little variation in communities between seasons. The proportion of sites occupied by taxa declined with increasing niche position, and decreasing niche breadth. Occupancy was predicted well by a combination of these two factors (niche position and breadth), but not mean local abundance, as the abundance-occupancy link was an upper-limit unimodal relationship. On average, community structure was linked more strongly to environmental variables in the wet season. Our findings demonstrate the clear role of spatial, but not temporal, turnover in assemblages, which likely reflects the environmental heterogeneity of this region. This is further supported by the fact that regional occupancy was mostly related to niche characteristics, particularly niche position. We emphasise the importance of continued basic and applied ecological work in this important biogeographic region to enable better protection of its biodiversity.

Drivers of macroinvertebrate community structure in unmodified streams

Often simple metrics are used to summarise complex patterns in stream benthic ecology, thus it is important to understand how well these metrics can explain the finer-scale underlying environmental variation often hidden by coarser-scale influences. I sampled 47 relatively pristine streams in the central North Island of New Zealand in 2007 and (1) evaluated the local-scale drivers of macroinvertebrate community structure as well as both diversity and biomonitoring metrics in this unmodified landscape, and (2) assessed whether these drivers were similar for commonly used univariate metrics and multivariate structure. The drivers of community metrics and multivariate structure were largely similar, with % canopy cover and resource supply metrics the most commonly identified environmental drivers in these pristine streams. For an area with little to no anthropogenic influence, substantial variation was explained in the macroinvertebrate community (up to 70% on the first two components of a partial least squares regression), with both uni- and multivariate approaches. This research highlights two important points: (1) the importance of considering natural underlying environmental variation when assessing the response to coarse environmental gradients, and (2) the importance of considering canopy cover presence when assessing the impact of stressors on stream macroinvertebrate communities.

Environmental Controls on River Assemblages at the Regional Scale: An Application of the Elements of Metacommunity Structure Framework

Understanding factors that structure regional biodiversity is important for linking ecological and biogeographic processes. Our objective was to explore regional patterns in riverine benthic invertebrate assemblages in relation to their broad positioning along the river network and examine differences in composition, biodiversity (alpha and beta diversity), and environmental drivers. We up-scaled methods used to examine patterns in metacommunity structure (Elements of Metacommunity Structure framework) to examine faunal distribution patterns at the regional extent for 168 low-mountain stream invertebrate assemblages in central Germany. We then identified the most influential environmental factors using boosted regression trees. Faunal composition patterns were compartmentalised (Clementsian or quasi-Clementsian), with little difference from headwaters to large rivers, potentially reflecting the regional scale of the study, by crossing major catchment boundaries and incorporating different species pools. While idealised structures did not vary, environmental drivers of composition varied considerably between river sections and with alpha diversity. Prediction was substantially weaker, and the importance of space was greater, in large rivers compared to other sections suggesting a weakening in species sorting downstream. Further, there was a stronger transition in composition than for alpha diversity downstream. The stronger links with regional faunal composition than with richness further emphasises the importance of considering the alternative ways in which anthropogenic stressors are operating to affect biodiversity patterns. Our approach allowed bridging the gap between local (or metacommunity) and regional scales, providing key insights into drivers of regional biodiversity patterns.

Climatic and Catchment-Scale Predictors of Chinese Stream Insect Richness Differ between Taxonomic Groups

Little work has been done on large-scale patterns of stream insect richness in China. We explored the influence of climatic and catchment-scale factors on stream insect (Ephemeroptera, Plecoptera, Trichoptera; EPT) richness across mid-latitude China. We assessed the predictive ability of climatic, catchment land cover and physical structure variables on genus richness of EPT, both individually and combined, in 80 mid-latitude Chinese streams, spanning a 3899-m altitudinal gradient. We performed analyses using boosted regression trees and explored the nature of their influence on richness patterns. The relative importance of climate, land cover, and physical factors on stream insect richness varied considerably between the three orders, and while important for Ephemeroptera and Plecoptera, latitude did not improve model fit for any of the groups. EPT richness was linked with areas comprising high forest cover, elevation and slope, large catchments and low temperatures. Ephemeroptera favoured areas with high forest cover, medium-to-large catchment sizes, high temperature seasonality, and low potential evapotranspiration. Plecoptera richness was linked with low temperature seasonality and annual mean, and high slope, elevation and warm-season rainfall. Finally, Trichoptera favoured high elevation areas, with high forest cover, and low mean annual temperature, seasonality and aridity. Our findings highlight the variable role that catchment land cover, physical properties and climatic influences have on stream insect richness. This is one of the first studies of its kind in Chinese streams, thus we set the scene for more in-depth assessments of stream insect richness across broader spatial scales in China, but stress the importance of improving data availability and consistency through time.