Water is an important dispersal vector for seeds of riparian plants, but little is known on how this form of dispersal, called hydrochory, varies throughout seasons and through geomorphically different areas. Therefore, we trapped seeds in 25 x 25 Astroturf mats along a free-flowing boreal river for an entire year. Sites mirrored the full range of geomorphic variation in the area and traps were emptied before and after the annual peak flow. The traps collected 2,062 seeds before the spring flood and 16,157 during the spring flood. While most of the seeds were deposited in the summer-low water traps (from now on "water edge") before the spring flood, they were distributed more equally in the riparian traps (defined as the level 40 cm above the water edge; from now on "riparian zone") during the spring flood. We found 77 species in total, of which 71 were at least recorded during the spring flood. Species numbers were higher at the water edge than at the riparian zone before and during the spring flood. Only 6.6% of the seeds, from 26 species, were viable. Most viable seeds were found in the water edge traps and, during the spring flood, in the riparian zone traps. While species that are primarily dispersed by water were not the most abundant among the viable seeds, they were the most species-rich group. This means that hydrochory is an important dispersal mechanism for riparian vegetation and an important contributor to riparian plant diversity in free-flowing rivers, and that changes in flow regimes, such as by flow regulation, can affect riparian vegetation composition.

Non-technical summary There has been a long history of conflicts, studies, and debate over how to both protect rivers and develop them sustainably. With a pause in new developments caused by the global pandemic, anticipated further implementation of the Paris Agreement and high-level global climate and biodiversity meetings in 2021, now is an opportune moment to consider the current trajectory of development and policy options for reconciling dams with freshwater system health. Technical summary We calculate potential loss of free-flowing rivers (FFRs) if proposed hydropower projects are built globally. Over 260,000 km of rivers, including Amazon, Congo, Irrawaddy, and Salween mainstem rivers, would lose free-flowing status if all dams were built. We propose a set of tested and proven solutions to navigate trade-offs associated with river conservation and dam development. These solution pathways are framed within the mitigation hierarchy and include (1) avoidance through either formal river protection or through exploration of alternative development options; (2) minimization of impacts through strategic or system-scale planning or re-regulation of downstream flows; (3) restoration of rivers through dam removal; and (4) mitigation of dam impacts through biodiversity offsets that include restoration and protection of FFRs. A series of examples illustrate how avoiding or reducing impacts on rivers is possible – particularly when implemented at a system scale – and can be achieved while maintaining or expanding benefits for climate resilience, water, food, and energy security. Social media summary Policy solutions and development pathways exist to navigate trade-offs to meet climate resilience, water, food, and energy security goals while safeguarding FFRs.

Ecosystem functioning of habitats at land-water interfaces, such as riparian forests and intertidal salt marshes or mangroves is predominantly driven by inundation. Whereas seasonality of ecological processes (i.e. phenology) and of hydrological extremes/events have been relatively well studied independently from each other their interdependence remains largely unknown. Filling this knowledge gap may become especially important in a changing climate as the timing of ecological and abiotic processes is already changing, often independently from each other. As these ecosystems are increasingly praised as Nature-based Solutions, predicting the ecosystem functioning of riparian forests and coastal wetlands under future climate change is crucial. Here, we will highlight the importance of match and mismatch of ecological and hydrological processes through a range of experiments and field observations in coastal wetlands from the single seedling to the ecosystem level. For riparian floodplains of Europe, we will show how the temporal relationships between flooding and thermal growing season have already changed in past decades, with currently unknown consequences. Finally, we will showcase methodological advances in field monitoring to better study these timing effects and offer conceptual insights to identify tipping points of ecosystem change along land-water interfaces. This presentation will focus on UPH ‘interfaces’ and ‘variability’.

Ecological restoration is becoming an increasingly common management tool worldwide. However, a challenge still exists on how to effectively monitor restoration outcomes and evaluate restoration success for ecological restoration managers. In this review, the goal is to evaluate whether the research in a degraded area has been sufficient for fostering efficient restoration measures and follow‐up of restoration success based on the Society for Ecological Restoration (SER) criteria. We selected the Inner Mongolian Steppe (IMS) in China as a model system. This area has been the subject of substantial research over the most recent years to understand degradation processes and restoration outcomes. We put together the variables used to assess degradation and restoration needs in the IMS and analyzed restoration results based on SER's nine criteria for evaluating restoration success. We found that the accomplished research in the IMS only partially supplied the data needed for evaluation of restoration success. The available results were sufficient for a proper evaluation of species composition and tentatively supported assessments of another seven criteria but not self‐sustainability. Grazing exclusion led to the fastest and most successful recovery of degraded steppe, but landscape‐scale processes during restoration in the IMS are still incompletely known. Our review supports large‐scale restoration of the IMS and emphasizes the need for long‐time monitoring for a more complete evaluation of the outcome of the IMS restoration following all SER's criteria.

Hydropeaking, defined as frequent and rapid variation in flow in regulated rivers with hydropower plants over a short period of time, usually sub-daily to weekly, alters hydraulic parameters such as water levels or flow velocity and exerts strong impacts on fluvial ecosystems. We evaluated the effects of hydropeaking on riverbank vegetation, specifically assessing the germination and establishment of seedlings and cuttings of plant species representing a variation in traits. We used seeds and seedlings and cuttings varying in size as phytometers, and transplanted them to riverbanks both above and below dams used for hydropower production in northern Sweden, selected to represent a gradient in hydropeaking intensity, and along a free-flowing reach. We also analyzed sub-daily water-level variables modified by hydropeaking to identify variables key in explaining the observed vegetation patterns. We found that plant responses to hydropeaking varied with species, with flood-intolerant species being the most strongly affected, as early as the germination stage. In contrast, seeds of flood-tolerant species managed to germinate and survive the early establishment phase, although strong erosive processes triggered by hydropeaking eventually caused most of them to fail. The fate of flood-intolerant species identifies germination as the most critical life-history stage. The depth and frequency of the inundation were the leading variables explaining plant responses, while the duration of shallow inundation explained little of the variation. The rise and fall rates of water levels were key in explaining variation in germination success. Based on the results, we propose restoration measures to enhance establishment of riparian plant communities while minimizing the impact on hydropower electricity production. Given the strong decrease in the germination of species intolerant to prolonged flooding with hydropeaking, planting of seedlings, preferably of large sizes, together with restrictions in the operation of the power plant during the establishment phase to enhance survival would be the best restoration option. Given the high probability of plant uprooting with hydropeaking, bank protection measures have the potential to increase riparian plant survival of all species, including flooding-tolerant species.

Although river restoration has increased rapidly, observations of successful ecological recovery are rare, mostly due to a discrepancy in the spatial scale of the impact and the restoration. Rivers and their ecological communities are a product of four river facets-hydrology, geomorphology, ecology and biogeochemistry-that act and interact on several spatial scales, from the sub-reach to the reach and catchment scales. The four river facets usually affect one another in predictable pathways (e.g., hydrology commonly controls geomorphology), but we show that the order in which they affect each other and can be restored varies depending on ecoregion and hydroclimatic regime. Similarly, processes at different spatial scales can be nested or independent of those at larger scales. Although some restoration practices are dependent of those at higher scales, other reach-scale restoration efforts are independent and can be carried out prior to or concurrently with larger-scale restoration. We introduce a checklist using the four river facets to prioritize restoration at three spatial scales in order to have the largest positive effect on the entire catchment. We apply this checklist to two contrasting regions-in northern Sweden and in southern Brazil-with different anthropogenic effects and interactions between facets and scales. In the case of nested processes that are dependent on larger spatial scales, reach-scale restoration in the absence of restoration of catchment-scale processes can frankly be a waste of money, providing little ecological return. However, depending on the scale-interdependence of processes of the river facets, restoration at smaller scales may be sufficient. This means that the most appropriate government agency should be assigned (i.e., national vs. county) to most effectively oversee river restoration at the appropriate scale; however, this first requires a catchment-scale analysis of feedbacks between facets and spatial scale interdependence.

River regulation may filter out riparian plants often resulting in reduced functional diversity, i.e., in the range of functions that organisms have in communities and ecosystems. There is, however, little empirical evidence about the magnitude of such reductions in different regions. We investigated the functional diversity patterns of riparian woody vegetation to streamflow regulation in boreal Sweden and Mediterranean Portugal using nine plant functional traits and field data from 109 sampling sites. We evaluated changes in mean plant functional traits as well as in indices of multidimensional functional traits, i.e., functional richness (FRic) and functional redundancy (FRed) within regions and between free-flowing and regulated river reaches. We found that regulation significantly reduced functional diversity in Sweden but not in Portugal. In Sweden, the increased magnitude of variations in water flow and water level in summer, the prolonged duration of extreme hydrological events, the increased frequency of high-water pulses, and the rate of change in water conditions were the likely main drivers of functional diversity change. Small riparian plant species with tiny leaves, poorly lignified stems, and shallow root systems were consistently associated with regulated sites in the boreal region. In Portugal, the similar functional diversity values for free-flowing and regulated rivers likely stem from the smaller streamflow alterations by regulation combined with the species legacy adaptations to the Mediterranean natural hydrological regimes. We conclude that streamflow regulation may reduce the functional diversity of riparian woody vegetation, but the magnitude of these effects will vary depending on the adaptations of the local flora and the patterns of streamflow disturbances. Our study provides insights into functional diversity patterns of riparian woody vegetation affected by regulation in contrasting biomes and encourages further studies of the functional diversity thresholds for maintaining ecosystems.

The computational tool InSTHAn (indicators of short-term hydrological alteration) was developed to summarize data on subdaily stream flows or water levels into manageable, comprehensive and ecologically meaningful metrics, and to qualify and quantify their deviation from unaltered states. The pronunciation of the acronym refers to the recording interval of input data (i.e., instant). We compared InSTHAn with the tool COSH-Tool in a characterization of the subdaily flow variability of the Colorado River downstream from the Glen Canyon dam, and in an evaluation of the effects of the dam on this variability. Both tools captured the hydropeaking caused by a dam operation, but only InSTHAn quantified the alteration of key flow attributes, highlighting significant increases in the range of within-day flow variations and in their rates of change. This information is vital to evaluate the potential ecological consequences of the hydrological alteration, and whether they may be irreversible, making InSTHAn a key tool for river flow management.

Stream networks both integrate abiotic and biotic landscape processes and transect the landscape, connecting different ecosystems and geologies longitudinally. The stream network can be divided into three process domains, as zones with distinct geomorphic processes: rapids, slowflowing reaches, and lakes. Biotic recovery has been variable after stream restoration and it is therefore important to understand where in the catchment it is most beneficial to focus restoration and how the different process domains influence the restoration outcome. Along a stream network, potential for organism dispersal, usually by hydrochory for riparian plants, control riparian community organization. Thus, we wanted to determine whether differences in recovery of riparian vegetation after restoration are a function of seed dispersal or habitat conditions. Our main objective was therefore to predict how the local and upstream source of riparian vegetation influence the restoration outcome. Our study was located in the boreal region of northern Sweden in the Hjuksån catchment. Hjuksån is a tributary of the free-flowing Vindel River, which in turn is the largest tributary to the Ume River. We studied three major factors: dispersal, germination and establishment success of riparian vegetation. In consistence with previous studies that stagnant waterbodies, such as lakes and fens are efficient seed traps, our study indicate that lakes retain more seeds than rapids and slow-flowing reaches, which will influence the riparian community recovery as less species will continue to downstream restored reaches. However, while the germination experiment showed that lakes had the highest germination success there were no such indications for the establishment. The higher germination success might partly be explained by lakes having a higher soil moisture then rapids, which is important for the germination success. Overall, this study indicated that the dispersal, germination and establishment is very low in naturally disconnected stream networks in northern Sweden and further restoration effort might be needed to aid the slow recovery.

Although river restoration has increased rapidly, observations of successful ecological recovery are rare, mostly due to a discrepancy in the spatial scale of the impact and the restoration. Rivers and their ecological communities are a product of four river facets-hydrology, geomorphology, ecology and biogeochemistry-that act and interact on several spatial scales, from the sub-reach to the reach and catchment scales. The four river facets usually affect one another in predictable pathways (e.g., hydrology commonly controls geomorphology), but we show that the order in which they affect each other and can be restored varies depending on ecoregion and hydroclimatic regime. Similarly, processes at different spatial scales can be nested or independent of those at larger scales. Although some restoration practices are dependent of those at higher scales, other reach-scale restoration efforts are independent and can be carried out prior to or concurrently with larger-scale restoration. We introduce a checklist using the four river facets to prioritize restoration at three spatial scales in order to have the largest positive effect on the entire catchment. We apply this checklist to two contrasting regions-in northern Sweden and in southern Brazil-with different anthropogenic effects and interactions between facets and scales. In the case of nested processes that are dependent on larger spatial scales, reach-scale restoration in the absence of restoration of catchment-scale processes can frankly be a waste of money, providing little ecological return. However, depending on the scale-interdependence of processes of the river facets, restoration at smaller scales may be sufficient. This means that the most appropriate government agency should be assigned (i.e., national vs. county) to most effectively oversee river restoration at the appropriate scale; however, this first requires a catchment-scale analysis of feedbacks between facets and spatial scale interdependence.

The Vindel River (Vindelfilven), Sweden, is 450 km long with a mean annual discharge of 190 m3/s and runs through sparsely populated areas in northern Sweden, joining the Ume River near the Baltic Sea. A severe decline in Atlantic Salmon Salmo salar and Brown Trout S. trutta during the past century was caused by (1) intense timber-floating activities starting in the mid-1800s, which degraded stream habitat; (2) hydropower development in the early 1900s, which limited or blocked upstream migration; and (3) an expanded coastal/ocean fishery after 1950, which overharvested anadromous populations. Restoration efforts during the 1970s and 1980s failed due to low efficacy of stocking programs and insufficient habitat restoration. A legislative reform to unify the fishery right owners (FROs) along the river helped initiate restoration efforts in the late 1990s, focusing on improving fish migration past a hydropower station and restoring degraded habitat. Sweden's membership in the European Union made large funding for restoration projects possible. The number of returning Atlantic Salmon increased significantly after migration conditions improved around the hydropower station and with stricter regulations on the offshore fishery. Successful habitat restoration was based upon gaining trust from landowners and FROs via extensive communication to gain access to their land. Restoration work was adaptive and experiences gained were incorporated into restoration guidelines. Involvement of universities, as a provider of expertise and as an unbiased interpreter of data, provided support to legal processes and when evaluating restoration measures. During the course of the restoration work, managers learned that stocking often did not produce satisfactory results. By studying historical documents from the timber-floating era, managers learned that the scope of modifications of the tributaries had been much greater than previously thought and that habitat restoration needed to be extensive. In many tributaries, the number of juvenile Brown Trout increased significantly after habitat restoration, sometimes dramatically exceeding expectations, which made the managers question the validity of established production estimates for northern boreal streams. The experience and knowledge gained from the Vindel River restoration served as the catalyst for many other major restoration projects in rivers emptying into the Baltic Sea.

Climate-change projections suggest large changes in riverine flow regime, which will likely alter riparian communities. In northern Europe, forecasts propose lower annual spring flood peaks and higher winter flows, resulting in narrower riparian zones. To estimate the impact of climate change on habitat extent of riparian plants, we developed a framework estimating the sensitivity and exposure of individual species to streamflow change, and surveyed five reaches along the free-flowing Vindel River in northern Sweden. We modeled the hydrologic niche of riparian plant species based on the probability of occurrence along gradients of flood frequency and duration and used predicted future water-level fluctuations (based on climate models and IPCC emission scenarios) to calculate changes in flow-related habitat availability of individual species. Despite projected increases in runoff, we predict most species to decrease in riparian elevational extent by on average 12-29% until the end of the century, depending on scenario. Species growing in the upper, spring-flood-controlled part of the riparian zone will likely lose most habitat, with the largest reductions in species with narrow ranges of inundation duration tolerance (decreases of up to 54%). In contrast, the elevational extent of most amphibious species is predicted to increase, but conditions creating isoëtid vegetation will become rarer or disappear: isoëtid vegetation is presently found in areas where ice formed in the fall settles on the riverbank during the winter as water levels subside. Higher winter flows will make these conditions rare. We argue that our framework is useful to project the effects of hydrologic change caused by climate change as well as other stressors such as flow regulation also in other regions. With few rivers remaining unaffected by dams and other human stressors, these results call for monitoring to detect species declines. Management to alleviate species losses might include mitigation of habitat degradation from land-use activities, more environmentally friendly flow schemes, and more intensive management options such as mowing riparian meadows no longer regularly maintained by recurrent floods.

The configuration of channels in stream networks is vital for their connectivity, biodiversity, and metacommunity dynamics. We compared the capacity of three process domains-lakes, slow-flowing reaches, and rapids-to disperse and retain plant propagules by releasing small wooden cubes as propagule mimics during the spring flood and recording their final locations. We also measured the geomorphic characteristics (planform, longitudinal profile, cross-sectional morphology, and wood) of each process domain. The three process domains all differed in morphology and hydraulics, and those characteristics were important in shaping the transport capacity of mimics. On average, lakes retained more mimics than slow-flowing reaches but did not differ from the retainment of rapids. Living macrophytes were the most efficient element trapping mimics. In rapids and slow-flowing reaches, most trapped mimics remained floating, whereas in lakes, most mimics ended up on the banks. The decay curves of retention varied substantially among and within process domains. The results suggest that managers who rely on natural recovery of restored sites by means of plant immigration may benefit from understanding landscape patterns when deciding upon the location of restoration measures in stream networks.

We tested whether the recovery of riparian vegetation along rapids that have been restored after channelisation for timber floating can be predicted based on floristic and geomorphic characteristics of surrounding landscape units. Our study was located along tributary stream networks, naturally fragmented in rapids, slow-flowing reaches, and lakes (i.e. process domains), in the Vindel River catchment in northern Sweden. We tested whether landscape characteristics, specifically to what extent the geomorphology (affecting local abiotic conditions), species richness, and species composition (representing the species pool for recolonisation), as well as the proximity to various upstream process domains (determining the dispersal potential), can predict post-restoration recovery of riparian vegetation. Our results indicate that post-restoration recovery of riparian vegetation richness or composition is not strongly related to landscape-scale species pools in these streams. The restored rapids were most similar to upstream rapids, geomorphically and floristically, including plant traits. Species richness of adjacent landscape units (upstream process domains or lateral upland zone) did not correlate with that of restored rapids, and proximity of upstream rapids or other process domains was only weakly influential, thus diminishing support for the hypothesis that hydrochory or other means of propagule dispersal plays a strong role in riparian vegetation community organisation after restoration in this fragmented stream network. We conclude that, in these naturally fragmented stream systems with three discrete process domains (rapids, slow-flowing reaches and lakes), hydrochory is probably not the main predictor for short-term riparian vegetation recovery. Therefore, other factors than landscape context can serve in prioritising restoration and, in these systems, local factors are likely to outweigh landscape connectivity in the recovery of riparian vegetation.

Free-flowing rivers (FFRs) support diverse, complex and dynamic ecosystems globally, providing important societal and economic services. Infrastructure development threatens the ecosystem processes, biodiversity and services that these rivers support. Here we assess the connectivity status of 12 million kilometres of rivers globally and identify those that remain free-flowing in their entire length. Only 37 per cent of rivers longer than 1,000 kilometres remain free-flowing over their entire length and 23 per cent flow uninterrupted to the ocean. Very long FFRs are largely restricted to remote regions of the Arctic and of the Amazon and Congo basins. In densely populated areas only few very long rivers remain free-flowing, such as the Irrawaddy and Salween. Dams and reservoirs and their up- and downstream propagation of fragmentation and flow regulation are the leading contributors to the loss of river connectivity. By applying a new method to quantify riverine connectivity and map FFRs, we provide a foundation for concerted global and national strategies to maintain or restore them.

Climate change will have large consequences for flooding frequencies in freshwater systems. In interaction with anthropogenic activities (flow regulation, channel restoration and catchment land-use) this will both increase flooding and drought across the world. Like in many other ecosystems facing changed environmental conditions, it remains difficult to predict the rate and trajectory of vegetation responses to changed conditions. Given that critical ecosystem services (e.g. bank stabilization, carbon subsidies to aquatic communities or water purification) depend on riparian vegetation composition, it is important to understand how and how fast riparian vegetation responds to changing flooding regimes. We studied vegetation changes over 19 growing seasons in turfs that were transplanted in a full-factorial design between three riparian elevations with different flooding frequencies. We found that (a) some transplanted communities may have developed into an alternative stable state and were still different from the target community, and (b) pathways of vegetation change were highly directional but alternative trajectories did occur, (c) changes were rather linear but faster when flooding frequencies increased than when they decreased, and (d) we observed fastest changes in turfs when proxies for mortality and colonization were highest. These results provide rare examples of alternative transient trajectories and stable states under field conditions, which is an important step towards understanding their drivers and their frequency in a changing world.

An Amendment to this paper has been published and can be accessed via a link at the top of the paper.

y In a changing climate, time sensitive ecological interactions such as pollination and predation are vulnerable to temporal mismatch with direct consequences for ecosystem functioning. It is not known if synchrony and asynchrony of ecological and physical processes such as flood disturbance and plant phenology may similarly be affected by climate change. Here, by spatially merging temperature and flood peak data, we show for the first time that in Central and Eastern Europe, annual river flood peaks increasingly occur within the thermal growing season. This is due to the combined effect of earlier spring onsets and later flood peaks. Such increased physical-phenological synchrony may especially impact river biogeomorphology and riparian floodplain ecosystem functioning through uprooting of seedlings and increased hydraulic roughness during major flood events.

Aims Many rivers across the globe are severely impacted by changed flooding regimes, resulting in drastic shifts in vegetation, but the processes driving the exchange of flood-sensitive and flood-tolerant species are understood less. We studied the role of long-term and recent flooding histories for riparian plant recruitment in response to various changes in flooding regime. Location Vindel River catchment (Northern Sweden). Methods We experimentally changed long-term flooding regimes by transplanting turfs between high and low elevations in 2000 and in 2014 (n = 8 per treatment). We sowed seeds of five riparian species in both transplanted turfs and non-transplanted controls and counted seedling numbers over two growing seasons. Further, we inventoried natural seedling frequencies in 190 plots in 19 reaches in 2013 and 2014, and related natural seedling numbers to plot flooding history in the period 2012-2014. Results We observed effects of long-term flooding history in the second year of the transplantation study (2015), but not in the first year. In 2015, turfs transplanted to locations with less flooding resulted in higher plant recruitment while transplantation to sites with more frequent flooding reduced recruitment compared to the controls. Since these differences were only found in recently transplanted turfs and not in older turfs, the legacy effect of long-term flooding history can be transient. In the field seedling survey, similar differences were found between flooding-history categories in 2013, but not in 2014, when the moisture conditions of the most recent year determined flooding. Further, lowest seedling numbers were observed when the previous flooding occurred in winter, and higher seedling numbers when floods occurred in spring or not at all. Conclusions Both long-term and recent flooding histories can affect plant recruitment, and their influence should be taken into account when designing restoration projects.

Although ecological restoration generally succeeds in increasing physical heterogeneity, many projects fail to enhance biota. Researchers have suggested several possible explanations, including insufficient restoration intensity, or time-lags in ecological responses that prevent detection of significant changes in short-term monitoring programs. This study aims to evaluate whether benthic macroinvertebrate communities responded to an expanded set of stream restoration measures within a study period of one to five years after completion of the restoration project. We studied 10 forest streams in northern Sweden that were channelized in the past for timber floating. Managers subjected six of these streams to habitat restoration, on each of these we selected two reaches, located in close proximity but differing in restoration intensity. In "basic" restored reaches, the restoration managers broke up the channelized banks and returned cobbles and small boulders to the main channel. In "enhanced" restoration reaches, they added additional large wood and boulders to reaches previously subjected to basic restoration, and rehabilitated gravel beds. The remaining four streams were not restored, and thus represent the baseline impacted (channelized) condition. We surveyed stream benthic assemblages before the enhanced restoration (year 2010) and three times afterward between 2011 and 2015. Five years after restoration, macroinvertebrate assemblages at the enhanced restored reaches were more differentiated from channelized conditions than those at basic-restored reaches. This reflected increased relative abundances of the insect orders Ephemeroptera and Trichoptera and the bivalve molluscs Sphaeriidae and decreased relative abundances of Chironomidae (Diptera). Analysis of functional traits provided further insights on the mechanistic explanations driving the recovery, e.g., indicating that the augmented channel retention capacity at enhanced restored reaches favored taxa adapted to slow flow conditions and more effectively retained passive aquatic dispersers. The increased restoration intensity in enhanced restored reaches has resulted in shifts in the composition of benthic macroinvertebrate assemblages, including increases in more sensitive taxa. These shifts became fully apparent five years after the enhanced restoration. Our results emphasize the value of longer-term monitoring to assess ecological responses following restoration, and of undertaking additional restoration as a valuable management option for previously restored sites that failed to achieve biotic recovery.