Tropical aquatic fungi

This paper reports on the distribution of fungal communities in aquatic habitats in tropical regions and highlights differences in the taxa observed in freshwater and marine habitats. Ascomycetes are dominant on substrata in marine environments, with few basidiomycetes and discomycetes. Equally, few freshwater basidiomycetes and discomycetes have been reported from the tropics. In marine habitats, Dothideomycetes dominate on mangrove substrata, and halosphaeriaceous species are most numerous on submerged woody substrata in coastal waters, while yeasts are common in seawater and estuarine habitats. In freshwater, Ingoldian anamorphic fungi are most numerous on decaying leaves, while ascomycetes (Dothideomycetes, Sordariomycetes) are prevalent on submerged/exposed woody substrata. Unique fungi are found in tropical waters and differ from those in temperate locations.     

Fungal biodiversity in aquatic habitats

Fungal biodiversity in freshwater, brackish and marine habitats was estimated based on reports in the literature. The taxonomic groups treated were those with species commonly found on submerged substrates in aquatic habitats: Ascomycetes (exclusive of yeasts), Basidiomycetes, Chytridiomycetes, and the non-fungal Saprolegniales in the Class Oomycetes. Based on presence/absence data for a large number and variety of aquatic habitats, about 3,000 fungal species and 138 saprolegnialean species have been reported from aquatic habitats. The greatest number of taxa comprise the Ascomycetes, including mitosporic taxa, and Chytridiomycetes. Taxa of Basidiomycetes are, for the most part, excluded from aquatic habitats. The greatest biodiversity for all groups occurs in temperate areas, followed by Asian tropical areas. This pattern may be an artifact of the location of most of the sampling effort. The least sampled geographic areas include Africa, Australia, China, South America and boreal and tropical regions worldwide. Some species overlap occurs among terrestrial and freshwater taxa but little species overlap occurs among freshwater and marine taxa. We predict that many species remain to be discovered in aquatic habitats given the few taxonomic specialists studying these fungi, the few substrate types studied intensively, and the vast geographical area not yet sampled.     

Global diversity of oribatids (Oribatida: Acari: Arachnida)

Oribatid mites are primarily terrestrial. Only about 90 species (less than 1% of all known oribatid species) from 10 genera are truly aquatic, with reproduction and all stages of their life cycle living in freshwater. Adaptation to aquatic conditions evolved independently in different taxa. However, many terrestrial species can also be found in aquatic habitats, either as chance stragglers from the surrounding habitats, or from periodic or unpredictable floodings, where they can survive for long periods. In spite of their low species richness aquatic oribatids can be very abundant in different freshwater habitats as in lentic (pools, lakes, water-filled microhabitats) or flowing waters (springs, rivers, streams), mainly on submerged plants. The heavily sclerotized exoskeletons of several species enables subfossil or fossil preservation in lakes or bog sediments. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Integrative freshwater ecology and biodiversity conservation

Freshwater ecosystems provide goods and services of critical importance to human societies, yet they are among the most heavily altered ecosystems with an overproportional loss of biodiversity. Major threats to freshwater biodiversity include overexploitation, water pollution, fragmentation, destruction or degradation of habitat, and invasions by non-native species. Alterations of natural flow regimes by man-made dams, land-use changes, river impoundments, and water abstraction often have profound impacts on lotic communities. An understanding of the functional interactions and processes in freshwater ecosystems presents a major challenge for scientists, but is crucial for effective and sustainable restoration. Most conservation approaches to date have considered single species or single level strategies. In contrast, the concept of ‘Integrative Freshwater Ecology and Biodiversity Conservation’ (IFEBC) proposed herein addresses the interactions between abiotic and biotic factors on different levels of organization qualitatively and quantitatively. It consequently results in a more holistic understanding of biodiversity functioning and management. Core questions include modeling of the processes in aquatic key habitats and their functionality based on the identification and quantification of factors which control the spatial and temporal distribution of biodiversity and productivity in aquatic ecosystems. The context and importance of research into IFEBC is illustrated using case studies from three major areas of research: (i) aquatic habitat quality and restoration ecology, (ii) the genetic and evolutionary potential of aquatic species, and (iii) the detection of stress and toxic effects in aquatic ecosystems using biomarkers. In conclusion, our understanding of the functioning of aquatic ecosystems and conservation management can greatly benefit from the methodological combination of molecular and ecological tools.     

DO FRESHWATER FISHES DIVERSIFY FASTER THAN MARINE FISHES? A TEST USING STATE‐DEPENDENT DIVERSIFICATION ANALYSES AND MOLECULAR PHYLOGENETICS OF NEW WORLD SILVERSIDES (ATHERINOPSIDAE)

Freshwater habitats make up only ～0.01% of available aquatic habitat and yet harbor 40% of all fish species, whereas marine habitats comprise >99% of available aquatic habitat and have only 60% of fish species. One possible explanation for this pattern is that diversification rates are higher in freshwater habitats than in marine habitats. We investigated diversification in marine and freshwater lineages in the New World silverside fish clade Menidiinae (Teleostei, Atherinopsidae). Using a time‐calibrated phylogeny and a state‐dependent speciation–extinction framework, we determined the frequency and timing of habitat transitions in Menidiinae and tested for differences in diversification parameters between marine and freshwater lineages. We found that Menidiinae is an ancestrally marine lineage that independently colonized freshwater habitats four times followed by three reversals to the marine environment. Our state‐dependent diversification analyses showed that freshwater lineages have higher speciation and extinction rates than marine lineages. Net diversification rates were higher (but not significant) in freshwater than marine environments. The marine lineage‐through time (LTT) plot shows constant accumulation, suggesting that ecological limits to clade growth have not slowed diversification in marine lineages. Freshwater lineages exhibited an upturn near the recent in their LTT plot, which is consistent with our estimates of high background extinction rates. All sequence data are currently being archived on Genbank and phylogenetic trees archived on Treebase.     

Global diversity of crayfish (Astacidae, Cambaridae, and Parastacidae––Decapoda) in freshwater

The freshwater crayfishes are distributed across all but the Indian and Antarctic continents with centers of diversity in the southeastern Appalachian Mountains in the Northern Hemisphere and in south–east Australia in the Southern Hemisphere. There are currently over 640 described species of freshwater crayfishes with an average of 5–10 species still being described each year. Freshwater crayfishes can serve as keystone species in aquatic habitats, but a few species are also significantly invasive and can cause impressive damage to the fragile freshwater habitat. Crayfishes inhabit caves, burrows, streams, lakes and strong burrowers can even be found in terrestrial habitats where they have burrowed to the water table or where rainfall is sufficiently abundant to provide the needed moisture. The freshwater crayfishes, like the habitats in which they are encountered, are generally endangered to some degree and conservation efforts would do well to focus on them as key elements of the freshwater ecosystem. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of lizards in freshwater (Reptilia: Lacertilia)

No lizards are strictly aquatic, but at least 73 species in 11 different families can be considered to regularly utilize freshwater habitats. There are no aquatic lizards in the Nearctic or Palearctic regions, whereas the Neotropics, Southeast Asia, and the Indo-Australian Archipelago support the greatest diversity of freshwater forms, particularly in the families Gymnophthalmidae, Scincidae and Varanidae. A number of larger aquatic lizards are harvested for food and for the reptile skin trade and several are CITES listed. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of mosquitoes (Insecta: Diptera: Culicidae) in freshwater

Mosquitoes that inhabit freshwater habitats play an important role in the ecological food chain, and many of them are vicious biters and transmitters of human and animal diseases. Relevant information about mosquitoes from various regions of the world are noted, including their morphology, taxonomy, habitats, species diversity, distribution, endemicity, phylogeny, and medical importance. Guest editors: E. V. Balian, C. Lévêque, H. Segers and K. Martens Freshwater Animal Diversity Assessment     

Global diversity of true bugs (Heteroptera; Insecta) in freshwater

The aquatic and semi-aquatic Heteroptera, consisting of the infraorders Leptopodomorpha, Gerromorpha, and Nepomorpha, comprise a significant component of the world’s aquatic insect biota. Within these three infraorders as a whole there are currently 23 families, 343 genera and 4,810 species group taxa considered valid, of which 20 families, 326 genera and 4,656 species inhabit freshwater. In addition, more than 1,100 unequivocally diagnosed species remain to be described. Aquatic Heteroptera occur on all continents except Antarctica, and are most numerous in the tropical regions, although there are many distinctly cold-adapted genera. Overall species richness is highest in the Neotropical and Oriental regions, which harbor 1,289 and 1,103 species, respectively. In comparison to these core tropical regions, species richness is significantly lower in the Afrotropical (799 species), Australasian (654 species), Palearctic (496 species), Nearctic (424 species) and Pacific (37 species) regions. Aquatic Heteroptera are notable for utilizing an exceptionally broad range of habitats, from marine and intertidal to arctic and high alpine, across a global altitudinal range of 0–4,700 m. Species may be found in almost every freshwater biotope, and many exhibit striking morphological adaptations to their aquatic environment, making them excellent subjects for ecological and biogeographic studies. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Evidence of local short‐distance spawning migration of tropical freshwater eels,and implications for the evolution of freshwater eel migration

Freshwater eels have fascinated biologists for centuries due to the spectacular long‐distance migrations between the eels’ freshwater habitats and their spawning areas far out in the ocean and the mysteries of their ecology. The spawning areas of Atlantic eels and Japanese eel were located far offshore in the Atlantic Ocean and the Pacific Ocean, respectively, and their reproduction took place thousands of kilometers away from their growth habitats. Phylogenetic studies have revealed that freshwater eels originated in the Indonesian region. However, remarkably little is known about the life histories of tropical freshwater eels despite the fact that tropical eels are key to understanding the nature of primitive forms of catadromous migration. This study found spawning‐condition tropical freshwater eels in Lake Poso, central Sulawesi, Indonesia, with considerably high gonadosomatic index values and with histologically fully developed gonads. This study provides the first evidence that under certain conditions, freshwater eels have conditions that are immediately able to spawn even in river downstream. The results suggest that, in contrast to the migrations made by the Atlantic and Japanese eels, freshwater eels originally migrated only short distances of <100 kilometers to local spawning areas adjacent to their freshwater growth habitats. Ancestral eels most likely underwent a catadromous migration from local short‐distance movements in tropical coastal waters to the long‐distance migrations characteristic of present‐day temperate eels, which has been well established as occurring in subtropical gyres in both hemispheres.     

Fungi in freshwaters: ecology, physiology and biochemical potential

Research on freshwater fungi has concentrated on their role in plant litter decomposition in streams. Higher fungi dominate over bacteria in terms of biomass, production and enzymatic substrate degradation. Microscopy-based studies suggest the prevalence of aquatic hyphomycetes, characterized by tetraradiate or sigmoid spores. Molecular studies have consistently demonstrated the presence of other fungal groups, whose contributions to decomposition are largely unknown. Molecular methods will allow quantification of these and other microorganisms. The ability of aquatic hyphomycetes to withstand or mitigate anthropogenic stresses is becoming increasingly important. Metal avoidance and tolerance in freshwater fungi implicate a sophisticated network of mechanisms involving external and intracellular detoxification. Examining adaptive responses under metal stress will unravel the dynamics of biochemical processes and their ecological consequences. Freshwater fungi can metabolize organic xenobiotics. For many such compounds, terrestrial fungal activity is characterized by cometabolic biotransformations involving initial attack by intracellular and extracellular oxidative enzymes, further metabolization of the primary oxidation products via conjugate formation and a considerable versatility as to the range of metabolized pollutants. The same capabilities occur in freshwater fungi. This suggests a largely ignored role of these organisms in attenuating pollutant loads in freshwaters and their potential use in environmental biotechnology.     

Latitudinal, habitat and substrate distribution patterns of freshwater ascomycetes in the Florida Peninsula

Freshwater ascomycetes are important decomposers of dead woody and herbaceous debris in aquatic habitats. Despite evidence of their ecological importance, latitudinal, habitat and substrate distributional patterns of freshwater ascomycetes are poorly understood. In this study, we examined the latitudinal and habitat distributional patterns, and substrate recurrences of freshwater ascomycetes by collecting dead submerged woody and herbaceous debris in lentic and lotic habitats at five selected sites along a north-central-south, temperate–subtropical latitudinal ecotone in Florida. One hundred and thirty-two fungal taxa were collected during the study. Seventy-four were meiosporic and 56 were mitosporic ascomycetes, while two species were basidiomycetes. Canonical analyses of principal coordinates (CAP) and Sørenson’s similarity index of species based on presence/absence data revealed a high turnover in species composition between the northern and southern sites, indicating a change in species composition along the temperate–subtropical latitudinal ecotone of the Florida Peninsula. Results from the ordination analysis indicated that freshwater ascomycete community composition is not significantly different between lentic and lotic habitats in Florida. The geographically broadly distributed species and species commonly found in Florida occurred in both habitats, whereas a number of new or rare species occurred in either lentic or lotic habitats, but not both. The same freshwater ascomycete species did not necessarily occur on both woody and herbaceous debris; of the 132 taxa collected, 100 were reported only on woody debris; 14 species occurred exclusively on herbaceous debris; and 18 species were found on both woody and herbaceous debris in lentic or lotic habitats. Implications of data from this study to the conservation and knowledge of biodiversity for freshwater ascomycetes is discussed.     

The structure of population genetic diversity in <Emphasis Type="Italic">Vallisneria</Emphasis><Emphasis Type="Italic">americana</Emphasis> in the Chesapeake Bay: implications for restoration

Submersed aquatic macrophyte beds provide important ecosystem services, yet their distribution and extent has declined worldwide in aquatic ecosystems. Effective restoration of these habitats will require, among other factors, reintroduction of genetically diverse source material that can withstand short- and long-term environmental fluctuations in environmental conditions. We examined patterns of genetic diversity in Vallisneria americana because it is a cosmopolitan freshwater submersed aquatic macrophyte and is commonly used for restoring freshwater habitats. We sampled 26 naturally occurring populations of V. americana in the Chesapeake Bay estuary and its tributaries and found that the majority of populations have high genotypic diversity and are not highly inbred. Fourteen of the populations had high allelic and genotypic diversity and could serve as source sites for restoration material. However, substantial geographic structuring of genetic diversity suggests that caution should be used in moving propagules to locations distant from their source. In particular, we suggest that propagules at least be limited within four primary geographic areas that correspond to freshwater tidal and non-tidal, oligohaline, and seasonally mesohaline areas of the Chesapeake Bay.     

Protected areas: A focus on Brazilian freshwater biodiversity

Brazil has a variety of aquatic ecosystems and rich freshwater biodiversity, but these components have been constantly damaged by the expansion of unsustainable activities. An array of different conservation strategies is needed, especially the creation of protected areas (PAs, hereafter). However, Brazil's PAs are biased towards terrestrial ecosystems and we argue that current PAs have limited efficacy in the protection of freshwater biodiversity. New PAs should better consider aquatic environments, covering entire basins, rivers and other freshwater habitats. We recommend ways to implement these PAs and provide guidance to avoid social impacts. Freshwater systems in Brazil provide essential goods and services but these ecosystems are being rapidly degraded and will be lost if not adequately protected.     

Evolutionary refugia and ecological refuges: key concepts for conserving Australian arid zone freshwater biodiversity under climate change

Refugia have been suggested as priority sites for conservation under climate change because of their ability to facilitate survival of biota under adverse conditions. Here, we review the likely role of refugial habitats in conserving freshwater biota in arid Australian aquatic systems where the major long‐term climatic influence has been aridification. We introduce a conceptual model that characterizes evolutionary refugia and ecological refuges based on our review of the attributes of aquatic habitats and freshwater taxa (fishes and aquatic invertebrates) in arid Australia. We also identify methods of recognizing likely future refugia and approaches to assessing the vulnerability of arid‐adapted freshwater biota to a warming and drying climate. Evolutionary refugia in arid areas are characterized as permanent, groundwater‐dependent habitats (subterranean aquifers and springs) supporting vicariant relicts and short‐range endemics. Ecological refuges can vary across space and time, depending on the dispersal abilities of aquatic taxa and the geographical proximity and hydrological connectivity of aquatic habitats. The most important are the perennial waterbodies (both groundwater and surface water fed) that support obligate aquatic organisms. These species will persist where suitable habitats are available and dispersal pathways are maintained. For very mobile species (invertebrates with an aerial dispersal phase) evolutionary refugia may also act as ecological refuges. Evolutionary refugia are likely future refugia because their water source (groundwater) is decoupled from local precipitation. However, their biota is extremely vulnerable to changes in local conditions because population extinction risks cannot be abated by the dispersal of individuals from other sites. Conservation planning must incorporate a high level of protection for aquifers that support refugial sites. Ecological refuges are vulnerable to changes in regional climate because they have little thermal or hydrological buffering. Accordingly, conservation planning must focus on maintaining meta‐population processes, especially through dynamic connectivity between aquatic habitats at a landscape scale.     

Does dispersal capacity matter for freshwater biodiversity under climate change?

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Global diversity of inland water cnidarians

Global diversity of inland water cnidarians is low, containing <40 species belonging to phylogenetically distinct groups representing independent invasion events: the common and cosmopolitan hydras (12–15 species); the sporadically occurring freshwater medusae (6–16 sp.); the Cordylophorinae (2 sp.); the parasitic Polypodium (1 sp.); the medusae occurring in saline lakes (4 sp.). Freshwater cnidarians inhabit nearly all types of freshwater on all continents (except Antarctica), but only a few species have cosmopolitan distributions. Due to uncertainty in species knowledge, fine scale regions of endemicity are not yet clear. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of fish (Pisces) in freshwater

The precise number of extant fish species remains to be determined. About 28,900 species were listed in FishBase in 2005, but some experts feel that the final total may be considerably higher. Freshwater fishes comprise until now almost 13,000 species (and 2,513 genera) (including only freshwater and strictly peripheral species), or about 15,000 if all species occurring from fresh to brackishwaters are included. Noteworthy is the fact that the estimated 13,000 strictly freshwater fish species live in lakes and rivers that cover only 1% of the earth’s surface, while the remaining 16,000 species live in salt water covering a full 70%. While freshwater species belong to some 170 families (or 207 if peripheral species are also considered), the bulk of species occur in a relatively few groups: the Characiformes, Cypriniformes, Siluriformes, and Gymnotiformes, the Perciformes (noteably the family Cichlidae), and the Cyprinodontiformes. Biogeographically the distribution of strictly freshwater species and genera are, respectively 4,035 species (705 genera) in the Neotropical region, 2,938 (390 genera) in the Afrotropical, 2,345 (440 genera) in the Oriental, 1,844 (380 genera) in the Palaearctic, 1,411 (298 genera) in the Nearctic, and 261 (94 genera) in the Australian. For each continent, the main characteristics of the ichthyofauna are briefly outlined. At this continental scale, ichthyologists have also attempted to identify ichthyological ‘‘provinces’’ that are regions with a distinctive evolutionary history and hence more or less characteristic biota at the species level. Ichthyoregions are currently identified in each continent, except for Asia. An exceptionally high faunal diversity occurs in ancient lakes, where one of the most noteworthy features is the existence of radiations of species that apparently result from intra-lacustrine speciation. Numerous fish-species flocks have been identified in various ancient lakes that are exceptional natural sites for the study of speciation. The major threats to fish biodiversity are intense and have been relatively well documented: overexploitation, flow modification, destruction of habitats, invasion by exotic species, pollution including the worldwide phenomena of eutrophication and sedimentation, all of which are interacting. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Latitudinal variation in global range-size of aquatic macrophyte species shows evidence for a Rapoport effect

1. To test hypotheses concerning the applicability of the Rapoport effect (RE: “species that occur at higher latitudes tend to have greater geographical range-size than species which have ranges limited to latitudes closer to the equator”) to aquatic macrophytes at global scale, we analysed the world latitudinal distribution and range-size of 1,083 vascular aquatic macrophyte species, from 91 genera in 11 families. We targeted macrophyte families strongly associated with inland aquatic habitats (i.e. with a zero, or only very low, proportion of constituent species which occur also in non-aquatic habitats), and which are distributed across a substantial latitudinal gradient, a necessary condition to test our hypotheses.
2. The macrophyte species present in these families include plants from all the normally accepted life form-defined functional groups of macrophytes, namely submerged, free-floating, floating-leaf rooted and emergent species, and represent the three major vascular taxonomic groups occurring as aquatic macrophytes (ferns/fern allies, monocots, and dicots). For the analysis, we used both latitude-only and areal measures of macrophyte species geographic range-size, within a 10 × 10° (latitude × longitude) grid of 238 grid cells, covering the six world ecozones (Palaearctic, Orient, Australasia, Nearctic, Neotropics, Afrotropics) that primarily contain inland freshwater and brackish macrophyte habitats.
3. The results provide new insight into the relationships between global range-size of macrophytes, latitude, and other potential spatio-environmental and anthropogenic drivers acting upon these plants at world scale. In particular, the outcomes indicated that: (1) the range-size versus latitude distribution of macrophytes shows evidence of a strong RE influence, with significantly greater species range-size at higher latitudes; and (2) the β-diversity pattern of species distribution along this latitudinal gradient is poorly explained by nestedness organisation, and species turnover is a more likely explanation of the observed changes in species distribution with latitude.
4. Spatio-environmental and anthropogenic variables other than latitude may also influence the observed global geographical pattern of macrophyte range-size, although their importance as predictors varies between individual families. Extent of agricultural land use, altitude, and historic (post-Quaternary) climate change velocity were all significant predictor variables for some families. However, interestingly, neither the area of land nor the area of waterbody present per grid cell were major influences on macrophyte range-size distribution.
5. Our finding of evidence for an RE, acting at global scale in aquatic macrophytes, contributes to increasing the generality of conclusions so far reached about the large-scale factors that drive patterns of species range-size at global scale. The study also provides a baseline for future macroecological work on aquatic plants, and potentially other freshwater organisms, particularly in the context of predicting how the world ranges of freshwater biota will respond to ongoing global environmental change.