Global diversity of ostracods (Ostracoda,Crustacea) in freshwater

There are close to 2,000 subjective species and about 200 genera of Recent non-marine Ostracoda. Together, Cyprididae (1,000 spp.) and Candonidae (c. 550 spp.) represent more than 75% of the extant specific diversity; the remaining 11 families comprise the other 25% of the species. The Palaearctic region has the highest absolute non-marine ostracod diversity, followed by the Afrotropical. The Australian region has the highest relative endemicity. About 90% of the species and 60% of the genera occur in one zoogeographical region only. This means that all the biological mechanisms which lead up to efficient dispersal and which are present in at least part of the non-marine Ostracoda (e.g. brooding, drought-resistant eggs, parthenogenesis) have not induced common cosmopolitan distributions in ostracods. Several habitats are hotspots for ostracod diversity and endemicity. For example, it appears that the ancient lakes hold up to 25% of the total ostracod diversity. Other speciation-prone habitats are groundwater, temporary pools and Australian salt lakes; in the latter two instances, cladogenesis has often been paralleled by gigantism. The present ostracod diversity results from 9 to 12 separate invasions of the non-marine habitat, starting about 400 Myr ago. Genetic diversity can be very different in different species, mostly, but not always, related to reproductive mode. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of tardigrades (Tardigrada) in freshwater

Tardigrada is a phylum closely allied with the arthropods. They are usually less than 0.5 mm in length, have four pairs of lobe-like legs and are either carnivorous or feed on plant material. Most of the 900+ described tardigrade species are limnoterrestrial and live in the thin film of water on the surface of moss, lichens, algae, and other plants and depend on water to remain active and complete their life cycle. In this review of 910 tardigrade species, only 62 species representing13 genera are truly aquatic and not found in limnoterrestrial habitats although many other genera contain limnoterrestrial species occasionally found in freshwater. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Global diversity of mayflies (Ephemeroptera, Insecta) in freshwater

The extant global Ephemeroptera fauna is represented by over 3,000 described species in 42 families and more than 400 genera. The highest generic diversity occurs in the Neotropics, with a correspondingly high species diversity, while the Palaearctic has the lowest generic diversity, but a high species diversity. Such distribution patterns may relate to how long evolutionary processes have been carrying on in isolation in a bioregion. Over an extended period, there may be extinction of species, but evolution of more genera. Dramatic extinction events such as the K-T mass extinction have affected current mayfly diversity and distribution. Climatic history plays an important role in the rate of speciation in an area, with regions which have been climatically stable over long periods having fewer species per genus, when compared to regions subjected to climatic stresses, such as glaciation. A total of 13 families are endemic to specific bioregions, with eight among them being monospecific. Most of these have restricted distributions which may be the result of them being the relict of a previously more diverse, but presently almost completely extinct family, or may be the consequence of vicariance events, resulting from evolution due to long-term isolation. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of mammals (Mammalia) in freshwater

Species that are dependant on, or adapted to, freshwater environments are found in almost all mammalian orders, and two orders, the Cetacea and the Sirenia, are strictly aquatic and include some freshwater-dependant species. Overall, the aquatic and freshwater-dependant species represent around 70 of the more than 1,200 living or recent genera of mammals, and occur in all continents except Antarctica. They include some of the most endangered species of mammals, and several have gone extinct or become critically endangered in recent decades. One of the main threats is habitat loss or degradation. This chapter provides an overview of the freshwater species within each order of mammals, their evolutionary history, their relations to humans and their conservation status. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Global diversity of nemerteans (Nemertea) in freshwater

Most ribborn worms (phylum Nemertea) are marine and only 22 of the currently named around 1,200 species are known from freshwater habitats (mainly lakes/ponds). They are all free-living benthic forms found in all continents except Antarctica. The vast majority of species have been recorded from the Palearctic region, but this may reflect sampling efforts rather than biogeography. Guest editors: E.V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of mysids (Crustacea-Mysida) in freshwater

In this article we present a biogeographical assessment of species diversity within the Mysida (Crustacea: Malacostraca: Peracarida) from inland waters. Inland species represent 6.7% (72 species) of mysid diversity. These species represent three of the four families within the Mysida (Lepidomysidae, Stygiomysidae, and Mysidae) and are concentrated in the Palaearctic and Neotropical regions. The inland mysid species distributional patterns can be explained by four main groups representing different freshwater invasion routes: (1) Subterranean Tethyan relicts (24 spp.); (2) Autochthonous Ponto-Caspian endemics (20 spp.); (3) Mysis spp. ‘Glacial Relicts’ (8 spp.); and (4) Euryhaline estuarine species (20 spp.). The center of inland mysid species diversity is the Ponto-Caspian region, containing 24 species, a large portion of which are the results of a radiation in the genus Paramysis. 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 and K. Martens Freshwater Animal Diversity Assessment     

Global diversity of aquatic macrophytes in freshwater

Aquatic macrophytes are aquatic photosynthetic organisms, large enough to see with the naked eye, that actively grow permanently or periodically submerged below, floating on, or growing up through the water surface. Aquatic macrophytes are represented in seven plant divisions: Cyanobacteria, Chlorophyta, Rhodophyta, Xanthophyta, Bryophyta, Pteridophyta and Spermatophyta. Species composition and distribution of aquatic macrophytes in the more primitive divisions are less well known than for the vascular macrophytes (Pteridophyta and Spermatophyta), which are represented by 33 orders and 88 families with about 2,614 species in c. 412 genera. These c. 2,614 aquatic species of Pteridophyta and Spermatophyta evolved from land plants and represent only a small fraction (∼1%) of the total number of vascular plants. Our analysis of the numbers and distribution of vascular macrophytes showed that whilst many species have broad ranges, species diversity is highest in the Neotropics, intermediate in the Oriental, Nearctic and Afrotropics, lower in the Palearctic and Australasia, lower again in the Pacific Oceanic Islands, and lowest in the Antarctic region. About 39% of the c. 412 genera containing aquatic vascular macrophytes are endemic to a single biogeographic region, with 61–64% of all aquatic vascular plant species found in the Afrotropics and Neotropics being endemic to those regions. Aquatic macrophytes play an important role in the structure and function of aquatic ecosystems and certain macrophyte species (e.g., rice) are cultivated for human consumption, yet several of the worst invasive weeds in the world are aquatic plants. Many of the threats to fresh waters (e.g., climate change, eutrophication) will result in reduced macrophyte diversity and will, in turn, threaten the faunal diversity of aquatic ecosystems and favour the establishment of exotic species, at the expense of native species. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of nematodes (Nematoda) in freshwater

Despite free-living nematodes being present in all types of limnetic habitats including unfavorable conditions that exclude many other meiobenthic invertebrates, they received less attention than marine and terrestrial forms. Two-fifths of the nematode families, one-fifth of the nearly 1800 genera and only 7% of the about 27,000 nominal species are recorded from freshwater habitats. The Dorylaimia are the most successful in freshwater habitats with nearly two-thirds of all known freshwater nematodes belonging to this subclass. Members of the subclass Enoplia are principally marine though include some exclusively freshwater taxa with extreme endemism. The subclass Chromadoria includes half of the freshwater nematode families and members of the Monhysterida and Plectida are among the most widely reported freshwater nematodes. Studies on freshwater nematodes show extreme regional bias; those from the southern hemisphere are extremely underrepresented, especially compared to European freshwater bodies. The majority of records are from a single biogeographic region. Discussion on nematode endemism is largely premature since apart from Lake Baikal, the nematofauna of ancient lakes as centers of speciation is limited and recent discoveries show high nematode abundance and diversity in cryptic freshwater bodies, underground calcrete formations and stromatolite pools potentially with a high number of new taxa. Guest editors: E.V. Balian, C. Lèvêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of copepods (Crustacea: Copepoda) in freshwater

The zoogeographic distributions of the 2,814 species of copepods reported from freshwater are analysed. Faunal diversity is compared between zoogeographic regions: the Palaearctic region has more than double the species richness of the next most diverse region, the Neotropical. Historical factors affecting levels of diversity are identified. More than 90% of all freshwater copepods are endemic to a single-zoogeographic region and endemic genera occur in all regions except Antarctica. Species that are not endemic to a single region include the highly vagile and cosmopolitan species occurring in four or more regions. The greatest faunal connectivity, as identified by Sørensen’s Index, is between Palaearctic and Nearctic regions, and identifies the Holarctic taxa. Key human-related issues, such as the role of copepods as vectors for human parasites and the losses caused by parasitic copepods in commercial aquaculture, are mentioned.     

Global diversity of cladocerans (Cladocera; Crustacea) in freshwater

Cladocera is a primarily-freshwater monophyletic group, an important component of the microcrustacean zooplankton. They inhabit most types of continental fresh and saline water habitats, occurring more abundantly in both temporary and permanent stagnant waters. Cladocera is an ancient group of Palaeozoic origin. About 620 species are currently known, but we estimate that the real number of species is 2–4 times higher. A number of currently-recognised widespread species can be expected to harbour extensive cryptic diversity. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of amphipods (Amphipoda; Crustacea) in freshwater

Amphipods are brooding peracaridan crustaceans whose young undergo direct development, with no independent larval dispersal stage. Most species are epibenthic, benthic, or subterranean. There are some 1,870 amphipod species and subspecies recognized from fresh or inland waters worldwide at the end of 2005. This accounts for 20% of the total known amphipod diversity. The actual diversity may still be several-fold. Amphipods are most abundant in cool and temperate environments; they are particularly diversified in subterranean environments and in running waters (fragmented habitats), and in temperate ancient lakes, but are notably rare in the tropics. Of the described freshwater taxa 70% are Palearctic, 13% Nearctic, 7% Neotropical, 6% Australasian and 3% Afrotropical. Approximately 45% of the taxa are subterranean; subterranean diversity is highest in the karst landscapes of Central and Southern Europe (e.g., Niphargidae), North America (Crangonyctidae), and Australia (Paramelitidae). The majority of Palearctic epigean amphipods are in the superfamily Gammaroidea, whereas talitroid amphipods (Hyalella) account for all Neotropic and much of the Nearctic epigean fauna. Major concentrations of endemic species diversity occur in Southern Europe, Lake Baikal, the Ponto-Caspian basin, Southern Australia (including Tasmania), and the south-eastern USA. Endemic family diversity is similarly centered in the Western Palearctic and Lake Baikal. Freshwater amphipods are greatly polyphyletic, continental invasions have taken place repeatedly in different time frames and regions of the world. In the recent decades, human mediated invasions of Ponto-Caspian amphipods have had great impacts on European fluvial ecosystems. Guest editors: E. V. Balian, C. Lévêque, H. Segers and K. Martens Freshwater Animal Diversity Assessment     

Global diversity of non-biting midges (Chironomidae; Insecta-Diptera) in freshwater

Chironomidae are common inhabitants of most aquatic habitats, and often dominate aquatic insect communities in both abundance and species richness. Species occur in all continents, including Antarctica, and most major oceanic islands that have been investigated. The family is divided into 11 subfamilies and 22 nominal tribes. Although individual species occur in a wide range of habitats from terrestrial to fully aquatic, a total of 339 genera and 4,147 species are unambiguously aquatic in their immature stages. Greatest species and generic richnesses occur in the Palaearctic Region and Nearctic Region, respectively, but this pattern may largely reflect historical patterns of past taxonomic research efforts. 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 gastropods (Gastropoda; Mollusca) in freshwater

The world’s gastropod fauna from continental waters comprises ∼4,000 valid described species and a minimum of 33–38 independent lineages of Recent Neritimorpha, Caenogastropoda and Heterobranchia (including the Pulmonata). The caenogastropod component dominates in terms of species richness and diversity of morphology, physiology, life and reproductive modes and has produced several highly speciose endemic radiations. Ancient oligotrophic lakes (e.g., Baikal, Ohrid, Tanganyika) are key hotspots of gastropod diversity; also noteworthy are a number of lower river basins (e.g., Congo, Mekong, Mobile Bay). But unlike many other invertebrates, small streams, springs and groundwater systems have produced the most speciose associations of freshwater gastropods. Despite their ecological importance in many aquatic ecosystems, understanding of even their systematics is discouragingly incomplete. The world’s freshwater gastropod fauna faces unprecedented threats from habitat loss and degradation and introduced fishes and other pests. Unsustainable use of ground water, landscape modification and stock damage are destroying many streams and springs in rural/pastoral areas, and pose the most significant threats to the large diversity of narrow range endemics in springs and ground water. Despite comprising only ∼5% of the world’s gastropod fauna, freshwater gastropods account for ∼20% of recorded mollusc extinctions. However, the status of the great majority of taxa is unknown, a situation that is exacerbated by a lack of experts and critical baseline data relating to distribution, abundance, basic life history, physiology, morphology and diet. Thus, the already considerable magnitude of extinction and high levels of threat indicated by the IUCN Red List of Threatened Species is certainly a significant underestimate. Guest editors: E. V. Balian, C. Lévêque, H. Segers and K. Martens Freshwater Animal Diversity Assessment     

Global diversity of Isopod crustaceans (Crustacea; Isopoda) in freshwater

The isopod crustaceans are diverse both morphologically and in described species numbers. Nearly 950 described species (∼9% of all isopods) live in continental waters, and possibly 1,400 species remain undescribed. The high frequency of cryptic species suggests that these figures are underestimates. Several major freshwater taxa have ancient biogeographic patterns dating from the division of the continents into Laurasia (Asellidae, Stenasellidae) and Gondwana (Phreatoicidea, Protojaniridae and Heterias). The suborder Asellota has the most described freshwater species, mostly in the families Asellidae and Stenasellidae. The suborder Phreatoicidea has the largest number of endemic genera. Other primary freshwater taxa have small numbers of described species, although more species are being discovered, especially in the southern hemisphere. The Oniscidea, although primarily terrestrial, has a small number of freshwater species. A diverse group of more derived isopods, the ‘Flabellifera’ sensu lato has regionally important species richness, such as in the Amazon River. These taxa are transitional between marine and freshwater realms and represent multiple colonisations of continental habitats. Most species of freshwater isopods species and many genera are narrow range endemics. This endemism ensures that human demand for fresh water will place these isopods at an increasing risk of extinction, as has already happened in a few documented cases. 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     

Global diversity of true and pygmy grasshoppers (Acridomorpha,Orthoptera) in freshwater

A small percentage of Orthoptera Acridomorpha is comprised species dependant on continental water ecosystems. However, as phytophagous insects, they are important at the basis of the trophic chain, mainly in regions with large permanent biota resulting from the pulses of the rivers. An assessment of the composition and origin of the populations from different biogeographical regions is attempted, and the state of our current knowledge is given in Tables 1, 2 and 3. 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     

Global diversity of crabs (Aeglidae: Anomura: Decapoda) in freshwater

The freshwater anomuran crabs of the family Aeglidae are all restricted to southern South America occurring in Chile, Brazil, Bolivia, Uruguay, Paraguay, and Argentina. The family consists of a single genus, Aegla, containing 63 currently described species. There are another 5–10 known yet undescribed species to complement this diversity. The aeglids occur in freshwater lakes, streams, rivers, and in caves with freshwater. The origin of the family appears to be from marine ancestors from the Pacific invading streams in Chile about 75 mya radiating both in Chile and again on the eastern side of the Andes, particularly in Brazil. Of the 63 species, 23 or 36.5% are considered under threat and are in need of conservation action. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment