dams is threatening linkages between the Andean headwaters and the lowland Amazon, with a huge river fragmentation [567], risking a start to irreversible processes, with drastic consequences on the conservation of biodiversity and for traditional and economic activities in the low lands [568,569]. A well-documented example of a dramatic impact on biodiversity and traditional activities is the construction (in 1994) of the Pak Mun Dam on the Mun river (Mekong basin), which caused the destruction of traditional fisheries and the decline of fish diversity. Roberts (2001) [570] identified the main impacts as being obstruction of breeding migrations, habitat conversion, and periodic dewatering or extreme flow variation downstream, combined with releases of oxygen-poor, warm, silty water from the reservoir. The situation is less critical in Africa but South-East Asia shows a worrying condition very similar to South America, with numerous built dams and dozens under construction or planned [571].

Another big threat for freshwater habitats is invasion by exotic species, recognized by now as the main driver of biodiversity change [572], particularly evident in lakes where non-indigenous species can rapidly become the dominant component of the ecosystem [573]. The introduction of non-native species is due to various causes, such as deliberate introduction to provide food or for sport purposes, for mosquito containment, or for ornamental purposes, or unwanted releases through the escape from aquaria, water gardens or aquacultures. An example of tremendous impact on biodiversity is Lake Victoria, where a single invader, the Nile perch (*Lates niloticus*), completely devastated the richest endemic fish fauna of the planet, without encountering predation or competition by native species [574,575]. Alien fishes that are not piscivores may also have large effects on their food: some authors [576] found a strong effect on aquatic invertebrate communities, via direct predation, by allochthonous Rainbow trout (*Oncorhynchus mykiss*) in high-Andean tropical streams. One of the most important classes of freshwater invaders includes filter-feeding molluscs, or those that feed on periphyton. These species can develop huge populations and consume so much primary production that they substantially influence the quantity and composition of primary producers. The interactions that radiate from primary producers can influence almost every part of the ecosystem [577]. The wetlands of southeastern Asia are invaded by the South American golden apple snail (*Pomacea canaliculata*), with severe pressure on the ecosystem through the elimination of macrophytes, causing increased concentrations of nutrients and large increases in phytoplankton [577]. The effects of herbivorous molluscs like the golden apple snail thus cause a shift in energy flow similar to that of severe eutrophication in shallow lakes [578]. Even exotic plants negatively affect ecosystems: an outstanding example is the water hyacinth (*Eichhornia crassipes*), a native of South America, but widespread in freshwater habitats of Africa and South-East Asia, and which is on the IUCN list of the 100 most dangerous invasive species. This species can occupy thousands of hectares of previously open water, forming extensive mats, heavily impacting the ecosystem: its rapid growth rate makes it capable of successfully competing with native plants and reducing light and oxygen resulting in a drastic change in submerged animal and plant communities. It can increase evapotranspiration, produce organic matter which increases the organic content of sediments, can favor the spread of diseases, and interferes with the use and management of water resources [579].

As mentioned before, alteration in land cover can affect sediment load but also directly impacts freshwater environments. Deforestation is particularly heavy in tropical America [562,580] and South-East Asia [581]: in Peninsular Malaysia, Sumatra, and Borneo, logging threatens peatlands at ever increasing levels, where, since 2007, industrial plantations have nearly doubled their extent [582]. So far, in these regions, managed land cover types have affected 50% of all peatlands: most of which are represented by oil palm plantation (73%), and the rest almost entirely occupied by plantations for paper pulp production, especially acacia [582]. While intact tropical peatlands can accumulate many meters of peat over thousands of years [541], degraded peatlands are susceptible to decomposition and fires [583]. This leads to soil subsidence [584], loss of biodiversity [585], and extremely high carbon dioxide emissions [586–588]. Finally, climate-change driven changes in temperature, total precipitation and rainfall pattern, besides increasing extreme climate events, can exacerbate all the consequences of threats outlined so far for tropical freshwater ecosystems. We can expect alterations in
water temperature, in hydrology and biogeochemical cycles, in evapotranspiration rates, and shifting species distribution, altering community structures and species interactions [15]. Different authors have explored the consequences of climate change on tropical freshwater habitats and their biodiversity but much work is still needed; among others, for predicted impacts in sub-Saharan wetlands [589], for impacts on Africa's freshwater biodiversity [590], for predicted impacts on the Brazilian Amazon wetlands [591], to investigated alterations in freshwater biodiversity in South-East Asia [592], and for impacts on tropical and subtropical Asian wetlands [593].

# 11. Arid-Climate Freshwater Habitats

Arid-climate freshwater habitats, including hot as well as high latitude cool- or cold-deserts are characterized by mean annual rainfall <250 mm, evaporation rates that exceed precipitation, and a variable hydrologic regime, with both droughts and flooding that vary in recurrence and severity from diel to century scales [594]. Temperature, humidity, and GW availability are among the main abiotic environmental factors influencing biodiversity and biogeography of arid freshwater groundwater, springs, streams, and lake ecosystems. However, a biocultural approach, including human population size and behavior, as has been applied in the study of oases in the Sahara and Sonoran Deserts, is needed to complement biological conservation strategies [154,595]. The casual observer might believe that arid regions support low species richness and limited biodiversity because of environmental harshness, but many taxa are well-represented in deserts, including aquatic, wetland, and riparian plants, microorganisms and aquatic faunas and assemblages (e.g., Figure 2A,J). High species richness in such habitats may be comparable to that in more mesic environments [596], and endemic diversity can be high. However, the diverse impacts of human pressures, including land-use, pollution, increased GW overexploitation, and global climatic change, pose dire threats for the sustainability of aridland-adapted freshwater species and habitats. Here we summarize arid-climate freshwater factors influencing ecology and biodiversity, with emphasis on microbial and algal ecology in desert springs, streams, and ciénegas (wet meadows).

GW and SW temperature and geochemistry strongly and non-linearly influence aridland springs biodiversity and ecology [115], and those in other freshwater ecosystems. Aridland stream nutrient dynamics are strongly influenced by SW-GW interactions, both within the stream channel and between the stream and adjacent riparian zones [597]. Nutrient transformations occur rapidly in channel floor sediments that are colonized by microbiota, and variable rates of processing result in spatial patchiness in nutrient availability in the water column. Nutrients and organic matter may accumulate along the channel margins or in riparian zones during dry periods and become remobilized during high flows, resulting in large temporal variability in nutrient dynamics with recolonization following such flows [598].

One of the most thoroughly-studied stream ecosystems in North America is Sycamore Creek in the Sonoran Desert, where warm stream temperature, high insolation and limited N availability all interact together to favour the proliferation of cyanobacterial mats, mainly including *Calothrix* sp. and *Anabaena* sp., often covering large portions of the stream bottom [599]. Freshwater stream-algal colonization following the flood succession usually starts with diatoms, then the filamentous green algae, and finally cyanobacterial mats [597]. This pattern can be related to N availability, as floods usually lead to a nitrogen pulse, thus alleviating N-limiting conditions for a brief time [600]. At early successional stages, algal standing stocks are low and mainly rely on nutrients from the water column and recycled from the sediments. As chlorophytes join the community and begin to form mats, leading to a decreased nutrient uptake from the environment and strong N limitation of this algal community, thus favoring colonization by cyanobacteria.

Freshwater organisms in arid-climate biomes have different strategic adaptations to GW and SW availability, temperature, geochemistry, and also to drying through behavior, physiology (e.g., pigment and lipid profiles), morphology, distribution (e.g., [601,602]), and biotic relationships. For example, many desert stream biota have the ability to rapidly recolonize after floods or rains and are distinctly
productive and diverse due to the high light availability and warmer temperatures. In another example, the frequency and duration of flooding have decreased in response to climatic change in arid Australian wetlands; however, waterbirds are facilitating recolonization of some invertebrates, plants (e.g., *Lemna*, *Typha*, and *Myriophyllum*) and charophytes (e.g., *Nitella*) through zoochorous propagule dispersal [603].

Harsh aridlands environments promote the evolution of endemism. For example, the warm, stenothermic, highly mineralized springs in Montezuma Well, Ash Meadows, Nevada, and Cuatro Ciénegas, Mexico, support high levels of endemic biodiversity [133,140]. Similarly, 67% of the metaphyton taxa in desert springs of the Bonneville Basin, Utah appear to have restricted distributions and are endemic to some individual springs [604]. That study also revealed dispersal limitations, with the absence of some species in more isolated springs. This observation supports Foissner's opinion (2006) [605] that algal distribution limitations exist, despite their potentially wide dispersal capacity. Dispersal limitations and adaptation similarly are responsible from much of the endemism among aridlands fauna (e.g., [606,607]).

Cantonati et al. (2015) [113] highlighted that the ambient springs in warm climate settings, including freshwater aridland ecosystems, host-diverse, species-rich cyanobacterial and algal assemblages, which, along with their adaptations to such habitats, are poorly known. The algal and cyanobacterial diversity of freshwater ponds and streams in the Central Death Valley Desert (Eastern California, USA) includes cyanoprokaryotes, green algae, and diatoms that dominate harsh thermophilic conditions there [608]. Morphotaxonomic surveys of freshwater ambient to slightly geothermal springs (ains; Figure 1AI) and drilled wells (birs) in the El-Farafra Oasis in the Western Desert of Egypt, revealed that those habitats often were impaired by human and livestock impacts (e.g., trampling and organic pollution) [609], as evidenced by the dominance by cyanoprokaryote bioindicators of eutrophic thermal freshwaters (*Geitlerinema splendidum*, *Jaaginema geminatum*, *J. subtilissimum*, *Kamptonema jasorvense*, *Limnothrix redekei*, *Oscillatoria animalis*, *O. limosa*, *O. princeps*, *O. tenuis*, and *Phormidium terebriforme*). Similarly, O₂-poor, warm springs in the Cuatro Ciénegas contain unusual cm-sized waterwarts built by an *Aphanothece*-like unicellular cyanobacterium and were suspended within a central, conically-shaped, 6-m deep well by upwelling waters, and supported a community of other epiphytic filamentous cyanobacteria and diatoms [610]. This unique jet-suspended, calcite-ballasted, colonial cyanobacterial community was narrowly adapted to conditions in that desert spring habitat. Also, Saber et al. (2018) [611] recently noticed unusual morphological characteristics in the streptophyte *Chara vulgaris* growing in the Springs of Moses in the Sinai Desert (Egypt). In both freshly-collected and cultured materials, thalli were delicate and the antheridia were shed early, the latter interpreted as an environmental adaptation to this highly-isolated and selective arid-climate freshwater ecosystem. Cyanoprokaryotes have wider biogeographic distributional niches than do other eukaryotic algal groups due to their different ecophysiological adaptive mechanisms to survive in arid-climate habitats and in addition to the production of several types of resting stages, which facilitate their environmental resistance and dispersal by wind or migrating animals [612].

As a prime example of a biodiverse aridland spring type, desert wetlands, or ciénegas, spanning the borderlands of Arizona (USA) and Sonora in México, are regions of high biodiversity conservation value. These environments contain an estimated 19% of species considered for federal protection in the region. Besides being crucial refugia for plants, mollusks and other invertebrates, native fish, amphibians, and reptiles, ciénegas also constitute a critical habitat for migratory birds. The increased and targeted habitat conservation of desert wetlands can yield great benefit to the maintenance of global biodiversity [154,613]. Similarly, the endemic biodiversity of Australian desert artesian springs includes 96 species and subspecies of plants, molluscs, crustaceans, and fishes, dominated by invertebrates with limited geographical ranges [614].

Environmental variability can significantly influence the distribution and abundance of endemic aridland freshwater biota, such as gastropods, particularly when extremes occur simultaneously over sustained time periods [615]. In general, freshwater desert springs are highly threatened ecosystems, but they often are remarkably constant environments that contain unique faunas with many endemic,
endangered, and cryptic aquatic taxa, like the endangered spring snails *Juturnia kosteri* (Figure 2AI) and *Pyrgulopsis roswellensis* found in springs in the Chihuahuan Desert (e.g., [616,617]), owing to their isolated, island-like nature and dispersal limitations [618]. Improved protection of these sensitive freshwater ecosystems is crucial for biodiversity conservation.

Information available on the biodiversity of arid-climate freshwater algae and cyanobacteria, as well as other freshwater springs-dependent taxa, is limited and understudied, particularly in North Africa and throughout South America. Much additional research is needed to fill these gaps in knowledge. Furthermore, the discovery of many endemic algal and cyanobacterial species in aridland freshwater ecosystems is predicted, especially as revealed through combined polyphasic approaches (i.e., morphotaxonomy, autecology, biochemistry, and genetics). For example, Saber et al. (2018) [611] recorded the new desmid species *Euastrum elfarafraense* (Figure 2AH) in an agricultural ditch fed by a typical inland-water rheocrenic hypothermal spring, "Ain El-Balad" in the El-Farafra Oasis, Western Desert of Egypt [619]. New and interesting cyanobacterial and green algal taxa have been recorded, but not yet described, from freshwater streams on bare rock inselbergs of *wadis* (Figure 1AJ) in the Eastern Desert of Egypt (Figure 1AH) (Saber A.A., unpublished data). Intensive human impacts on GW and SW, such as coarse-scale inputs of agricultural nutrients, release of domestic sewage and other pollutants, as well as the global climate change, are likely to cause severe structural and functional degradation of arid-climate freshwater habitats, with subsequent degradation of their biotic assemblages. Moreover, the resilience of these ecosystems from severe anthropogenic stress appears to be low, leading to a high risk of extirpation and extinction of many interesting and poorly-known rare and endemic species.

## 12. Freshwater Biodiversity Observation Network (FWBON)

The Freshwater Biodiversity Observation Network (FWBON) is a voluntary community of practice dedicated to tracking change in global freshwater biodiversity. It was established in 2016 as a biodiversity observation network affiliated with GEOBON (https://geobon.org/). FWBON promotes best practices for global freshwater biodiversity observations by helping to (1) improve the collection of harmonized data; (2) develop data standards and methodologies for data management and dissemination; (3) share biodiversity data across the world without compromising national concerns; (4) integrate biodiversity information with physical and chemical data; (5) generate products useful for sound management of rivers and their catchments, lakes, wetlands and subterranean aquatic ecosystems; and (6) integrate freshwater science and practice with terrestrial and coastal conservation objectives.

With members in over 50 countries, FWBON strives to bring freshwater biodiversity monitoring expertise and experience from across the world to global biodiversity observation platforms [620]. Its activities build on and support a multitude of projects and programs funded at local, national, and regional scales, e.g., the Circumpolar Freshwater Biodiversity Monitoring Program [418]. This enables FWBON to promote a global agenda that is firmly anchored in national and local capabilities and priorities. It also allows FWBON to facilitate the flow of knowledge on freshwater biodiversity both ways along a hierarchy of spatial scales from local to global. Many of FWBON's activities are connected to essential biodiversity variables (EBV; [621]). FWBON has played a leading role in promoting the implementation of the EBV concept in freshwater biodiversity monitoring across the world [622]. It has also helped in developing workflows to support the generation of EBVs from primary biodiversity data [623] and the definition and development of species population EBVs [624].

In focusing on tracking global change in freshwater biodiversity, FWBON is strongly driven by a freshwater biodiversity conservation agenda, and it collaborates with others who have a similar agenda. A recent example of such collaboration is the formation of the Alliance for Freshwater Life (AFL), which is a global initiative, uniting specialists in research, data synthesis, conservation, education and outreach, and policymaking [14].
## 13. Discussion

Effective conservation solutions are dependent on an understanding of hydrologic connectivity between different freshwater ecosystems (including connected coastal and marine systems). This was demonstrated recently in a spatial prioritization exercise in south-eastern Australia in which rivers, wetlands and aquifers were considered simultaneously, and this greatly improved the efficiency of regional conservation planning solutions [112]. However, considering hydrological connectivity alone is not sufficient to optimize conservation planning and get the best results from local conservation actions. Different aquatic ecosystems are also connected with one another and with terrestrial ecosystems through mechanisms not explained by hydrological connectivity [625,626]. Not considering these mechanisms might result in a failure to recognize important conservation solutions and opportunities. For example, terrestrial vegetation corridors may be critical for the persistence and dispersal of aquatic species if they have the function of protecting non-aquatic stages of these species and facilitating their dispersal [627]. Similarly, hydrological connectivity alone does not account for the input of organic matter into aquatic ecosystems. Hence, the information on the structure and spatial configuration of terrestrial vegetation could complement information on hydrological connectivity.

This integrated view (Figure 3), to be achieved, might benefit from the theoretical foundations provided by approaches grounded in ecohydrogeology [84], catchment geodiversity (rock type, soil type, and geomorphological richness) as a basis to infer freshwater biodiversity facets [435], and increased recognition that healthy, diverse freshwater ecosystems provide vital natural functions at the landscape scale (e.g., [628]).

Global biodiversity is challenged under the pressure of several human-induced changes of the global environment at an unprecedented rate. The extent and rate of this change is so relevant and so strongly linked to the exploitation of natural resources and to ecosystem processes that biodiversity change has to be considered an important global change in its own right [1]. Previous reviews [1,2,629,630] identified the following as the most important categories of drivers of changes in freshwater biodiversity and ecosystems at the global scale (Table 1): water pollution including eutrophication, flow modification, overexploitation, destruction or habitat degradation, biotic exchanges (invasive non-native species; e.g., [631]), changes in land use, atmospheric CO₂ concentration, and nitrogen deposition and acid rain. Stendera et al. (2012) [632] highlighted that hydrological modifications are one of the main stressors of freshwater biodiversity, while Cañedo-Argüelles et al. (2016) [633] pointed out the dire consequences of salinization.

Strayer and Dudgeon (2010) [629] discussed four important challenges for freshwater conservation: (1) Climate change, which will endanger both freshwater species and human uses of fresh water, driving engineering responses that will further threaten the freshwater biota; (2) because freshwater extinctions are already ongoing, freshwater conservationists must be prepared to act now to prevent further losses, and engage more effectively with other stakeholders; (3) the gap between freshwater ecology and conservation biology needs to be bridged; (4) freshwater sciences societies and journals need to improve their historically poor record in publishing important papers and influencing practice in conservation ecology.

Climate-change effects are typically superimposed upon those of multiple stressors, and climate change is thus often regarded as a threat multiplier [634]. We thus think it is of paramount importance to favor studies and initiatives to determine and quantify freshwater-ecosystem resistance, resilience, and recovery to global climate change (e.g., [634]), because this allows assessment of their strategic role (e.g., [635]) and value for biodiversity conservation. One possibility to increase resilience is artificial aquifer recharge (e.g., [636]), but its use to support freshwater biodiversity and ecological integrity is almost unexplored. A particular facet of climate change is also extreme events that are increasing in frequency and intensity, including ever stronger heat-waves and prolonged droughts. Segadelli et al. (2019) [47] in this VSI discuss a first attempt to predict and understand the ecological effects of extreme precipitation events in small montane catchments.
<figure><img src="image_8.png" /></figure>

**Figure 3.** Landscape perspective (schematic drawing) of the hydrologic cycle intended to focus on relationships and interactions among all freshwater ecosystem types dealt with in the present paper. Though generalized as much as possible, the depiction is placed in a specific (montane) setting for the sake of concreteness. Labels: GDEs = groundwater and dependent ecosystems, Sp = springs, SpSt = spring-fed streams, GlSt = glacial streams, St = streams, Ri = rivers, TeLa = tectonic lakes, HMLa = high-mountain lakes, ObLA = oxbow lakes, Re = reservoirs, Mi = mires, SWE = small wetland ecosystems.
In this paper, we aimed at connecting the different accounts of freshwater ecosystems presented by individual papers in this VSI. These accounts show how varied freshwater habitats are and describe current approaches to studying, monitoring, and assessing these disparate habitats. The paper also reveals major differences among experts working on different ecosystems, suggesting that the varied ways freshwater ecosystems are studied and assessed could be as much a reflection of the traditions followed by the specialists studying them and of the geographical and sociopolitical context in which they operate [9] as of physical and ecological differences among the ecosystems.

In the face of continuously shrinking resources both for freshwater-biodiversity inventories and environmental-quality assessments, the question arises whether freshwater-biodiversity inventories can be funded independently from assessment and monitoring efforts [637]. The decoupling of biodiversity studies from assessments is unfortunately most likely to cause a strong reduction in opportunities to acquire large-scale distribution data for many categories of organisms. This challenge is not unique to freshwaters. No matter how big their budget, research projects with short life-times and specific objectives cannot meet the data needs for large-scale assessments of the status of biodiversity and how this might be changing over years and decades. Hence, it is important to have a solid conceptual framework that helps to utilize data from multiple sources to fill gaps in the space-time-species space [624]. There is also a need for practical solutions, e.g., workflows [623], to convert disparate primary data into biodiversity variables that can be used to measure biodiversity change at spatial and temporal scales relevant to decision-making and policy development. Measuring the different dimensions of biodiversity in freshwaters at multiple spatial scales presents some specific challenges [622] which must be addressed in the near future.

Empirical, experimental, and theoretical research on species loss focused on the attempt to link biodiversity loss to a reduction in the capability of ecosystems to perform energy and matter cycling. Cardinale et al. (2006) [638] showed that the magnitude of these effects is ultimately determined by the identity of the species involved. They noted that a key challenge for future research is to detail more accurately how the traits that determine vulnerability to extinction are related to functional dominance in communities, and underlined that, until that time, a precautionary approach to preserving as much biodiversity as possible is justified. Biodiversity loss aroused not only concern but also interest and controversy among theoretical ecologists. The main cause of controversy was the uncertainty as to how results describing the relationship between species diversity and ecosystem processes, and identifying functionally important species, might scale up to landscape and regional levels, and generalize across ecosystem types [639]. Moreover, many recent experiments on the topic were performed using terrestrial plant-dominated systems (e.g., [640]), while the vast areas of biodiversity that involve small organisms (i.e., bacteria, archea, protists, microarthropods), which drive the bulk of ecosystem processes, are likely to be of particular importance [639]. From a strictly functional point of view, species matter so far as their individual traits and interactions contribute to maintaining the functioning and stability of ecosystems and biogeochemical cycles. However, there are a number of reasons, including cultural, economic, and aesthetic, justifying the need for biodiversity conservation (e.g., [639]). Freshwater biodiversity provides a broad variety of valuable goods and services for human societies, some of which are irreplaceable [641]. The value of this biodiversity has several components: (1) its direct contribution to economic productivity, (2) its "insurance" value in light of unexpected events, (3) its value as a storehouse of genetic information, and (4) its value in supporting the provision of ecosystem services (e.g., cleaning water) [2]. Estimates of the full value of biodiversity need to account for each of these four components. However, the precise impacts of biodiversity change will vary with the ecosystem type and with the processes and properties considered.

At smaller geographic scales, there is substantial species turnover (i.e., β-diversity [161]) among drainage basins and water bodies, and many freshwater species have restricted ranges (e.g., [642]). These attributes combine with endemism to produce a lack of "substitutability" among freshwater habitat units. This means that protection of one or a few water bodies cannot preserve all freshwater biodiversity within a region, or even a significant proportion of it [2].
While preservation of intact freshwater bodies and their biodiversity remains a priority, it is important to recognize the potential that partly degraded habitats may have to support significant portions of their original biodiversity [2]. In an attempt to move towards such “win-win” solutions, Rosenzweig (2003) [643] advocates an approach to enhancing species richness in human-dominated landscapes termed “reconciliation ecology”. It is to such strategies that freshwater scientists should consider turning, where appropriate, rather than persisting only in attempts to preserve intact ecosystems in the face of increasing human pressure [2]. On the other hand, it is very easy to compromise freshwaters, and often almost impossible to restore [644,645].

Terrestrial conservation strategies tend to emphasize areas of high habitat quality that can be protected. This “fortress conservation” is likely to be unsuccessful for freshwaters [646] and may even be counterproductive [647] for river segments or lakes in unprotected drainage basins unless the boundaries are drawn at a catchment scale, which is virtually never the case [2,648].

The particular vulnerability of freshwater biodiversity also reflects the fact that freshwater is a resource for humans that may be used, diverted, or contaminated in ways that compromise its value as a habitat for organisms [2]. In the vast majority of disagreements over multiple uses of water, whether they are international or on a local scale, the allocation of water to maintain aquatic biodiversity is largely disregarded [649]. Humans have appropriated more than half of renewable freshwater to their own use [650], but in spite of this, nearly 80% of the world population is highly threatened as concerns water security [651]. In exploiting this resource for different uses (e.g., urban, agricultural, industrial), the vital role of water in supporting ecosystems has been widely overlooked (e.g., [652]). To protect the world's freshwater resources, human and biodiversity perspectives on water security need to be jointly considered [651]. There has indeed been growing recognition that preserving ecological functionality and the biological complexity of freshwater ecosystems is the only way to safeguard many valuable services and benefits to society (e.g., [652]).

**Author Contributions:** M.C. conceived and designed the paper (including tables & figues). Authors revising and editing the whole paper: M.C., S.P., C.M.P., L.E.S., E.T., J.H., J.S.R., S.S. (Figure 3, technical editing). Section Lead Authors: M.C. (Glacial Streams, High-Mountain Lakes), L.E.S. (Springs), E.T. (Urban), J.H. (Boreal, Arctic), J.S.R. (Headwaters, Streams, Rivers), R.B. (Oxbow Lakes, SWE, lead for technical editing), A.B. (Tropical+ technical editing), N.C. (Med. Streams), M.Č. (Reservoirs), D.M.P.G. (Groundwater), M.H. (Mires), I.H. (Antarctic), Z.L. (Ancient Lakes), L.N.-F. (Mediterranean Pools, Reservoirs), A.A.S. (Arid). Section Co-Authors: M.C. (Springs, Large Lakes, Mires), C.M.P. (Streams, Rivers), M.D.C. (Groundwater, references, formatting), B.F. (Groundwater), P.B.H. (Ancient Lakes), J.K. (Reservoirs), P.Z. (Reservoirs), L.E.S. (Arid). All authors have read and agreed to the published version of the manuscript.

**Funding:** This research received no external funding.

**Acknowledgments:** Tamar Zohary (IOLR & SIL, comments on an early version & photos). Nicola Angeli (MUSE; Figures 1 and 2 layout), Sebastian Birk (Duisburg-Essen, literature suggestions for Large Rivers). M.H. acknowledges project number GX19-28491X. The ‘Groundwater’ section was supported by AQUALIFE LIFE12 BIO/IT/000231. M.Č., J.K. and P.Z. were supported from ERDF/ESF project No. CZ.02.1.01/0.0/0.0/16_025/0007417. L.E.S. thanks S.W. Carothers and M.E. Melray for supporting his work. Diatom part presented as plenary lecture at the North American Diatom Symposium organized by Kalina Manoylov & James Nienow (supporting M.C. for travel costs). We thank our supporting Institutions. Authors are also grateful for the very valuable suggestions provided by anonymous reviewers and the editor that helped to improve the original manuscript.

**Conflicts of Interest:** The authors declare no conflict of interest.

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