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     

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 stoneflies (Plecoptera; Insecta) in freshwater

Plecoptera, or stoneflies, is a small order of hemimetabolous insects: according to our data, more than 3,497 species have been described so far in the world. The total number of species has enormously increased in the last 30 years (2,000 species estimated in 1976) and, if the trend continues, then it will nearly double in the near future. The order is divided into the suborders Arctoperlaria and Antarctoperlaria, and includes 16 families: 12 arctoperlarian and 4 antarctoperlarian. The Arctoperlaria account for a total number of 3,179 species, and Antarctoperlaria, only 318 species. The total number of genera is 286. We give in this article the estimated number of species for each family. The fauna and diversity of stonefly in North America (650 species reported) and Europe (426 species) are best known. Nevertheless, in the last 25 years, a mean of 2.6 Plecoptera species per year were described in Europe. Stonefly-faunas of Australia (191 species, Tasmania included) and New Zealand (104 species) are relatively well-known, while our knowledge of the Plecoptera of Central and South America (95 and 378 species respectively) is poor and still not representative of the real diversity. Africa has a reduced stonefly fauna (126 species). Asian stonefly diversity (approximately 1,527 species) is much greater than that of Europe or North America despite the fact that, except for Japan and Asiatic Russia that have been well studied, our knowledge of the remaining Asiatic areas is extremely poor. Even though our data indicate the Holarctic Region as the diversity hot-spot for the order, the analysis of the specific diversity divided by family suggests also an important role of tropical stoneflies. Guest editors: E.V. Balian, C. Lévêque, H. Segers & 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 hymenopterans (Hymenoptera; Insecta) in freshwater

A summary of the known species of aquatic Hymenoptera is presented. In total, 150 species from 11 families are recognized as aquatic (0.13% of the total described species). This number is likely an underestimate, because of the high percentage of undescribed species and a lack of knowledge of host range and behaviour for most species. All aquatic Hymenoptera are parasitoids. Many species have relatively dense pubescence to trap air and elongate, tarsal claws to grip the substrate, when underwater. Most species are known from the Holarctic and Oriental regions, but this is likely an artefact caused by lack of knowledge of other regions of the world. Aquatic behaviour has evolved independently at least 50 times within the order. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Global diversity of craneflies (Insecta,Diptera: Tipulidea or Tipulidae <Emphasis Type="Italic">sensu lato</Emphasis>) in freshwater

The Tipulidae s.l.—craneflies—are one of the largest groups of the Diptera containing over 15,270 valid species and subspecies. The immatures of the majority of species live in aquatic or semiaquatic habitats. Some aquatic species live entirely submerged and lack functional spiracles, others come to the surface to take oxygen by using spiracles positioned at the end of the abdomen. Semiaquatic species occur in a wide range of habitats. The semiterrestrial and terrestrial larvae live in environments that are moist or at least humous. All adult craneflies are terrestrial. Conflicting hypotheses on the phylogenetic position of the Tipuloidea within the Diptera continue to exist: some authors consider them to represent one of the oldest lineages of the Diptera, others suppose a close relationship to the Brachycera, the true flies. Current systematic knowledge of the Tipuloidea indicates that the Palaearctic region contains the highest number of genus-group taxa, while the Neotropical region has the highest number of species and subspecies. The Afrotropical and Australasian regions are relatively poor respectively in genera and subgenera and in species and subspecies. The oldest fossils that represent the Tipuloidea date back to the Lower Triassic at about 240 million years. Present-day general distribution patterns of many higher taxa of Tipuloidea probably have a Pangean or Gondwanan origin. 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 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 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 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     

Species diversity and abundance of Tabanus spp. (Diptera: Tabanidae) in different habitats of Thailand

Tabanus spp. or horse flies (Diptera: Tabanidae) are haematophagous flies of medical and veterinary importance. They are known to cause trypanosomosis or surra in domestic and wild animals in Thailand. This study conducted an entomological survey of horse flies from different sites in Thailand. Horse flies were collected from three different habitats: primary forests, secondary forests and villages using Nzi traps between April 2012 and December 2016. A total of 1835 female horse flies were collected and 45 species were identified. The five most abundant species were T. striatus (25.45%), followed by T. megalops (21.36%), T. rubidus (14.82%), T. tamthaiorum (7.90%) and T. oxybeles (6.38%). The highest proportion of horse flies was collected in villages (39.13%), followed by primary forests (34%) and secondary forests (26.87%). The species diversity of horse flies in primary forests was higher than in other habitats. The results of this study may be used for a horse fly control program.     

Global diversity of butterflies (Lepidotera) in freshwater

In Lepidoptera, the subfamily Acentropinae and Pyraustinae of Crambidae (Pyraloidea) and the family Arctiidae (Noctuoidea) contain species with true aquatic larvae, which live submerged during larval development. In Pyraustinae and Arctiidae only a few species exhibit an aquatic life-history. From the latter, aquatic larvae are known from the Neotropical genus Paracles. The number of aquatic Paracles species is unknown. The Acentropinae are predominantly aquatic. They are distributed worldwide, and reach the highest diversity in tropical regions of South East Asia/Malesia and in the Neotropical Region. At present, the Acentropinae include a total of 50 genera and 737 described species. All genera, assigned to the subfamily, are listed in a table, and the numbers of included species are indicated. The taxonomy and phylogeney of the genera are inadequately known. The species have a minor economic importance, however, they are very sensitive to degradation of water quality and habitat destruction. Guest editors: E.V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

Chironomidae (Insecta: Diptera) from four protected areas in the state of Pernambuco,Brazil

New records and distributional notes of Chironomidae (Insecta, Diptera) are provided for four protected areas in the state of Pernambuco, northeastern Brazil. Additionally, we also present new records and update of distributional ranges from Brazil and the Neotropical Region. In total, 810 specimens belonging to 35 genera within the subfamilies Chironominae (22 taxa), Tanypodinae (11 taxa) and Orthocladiinae (2 taxa) were found. The subfamilies Chironominae and Tanypodinae predominated. Axarus Roback, 1980 Roback, S.S. (1980), ‘New name for Anceus Roback nec Anceus Risso’, Entomological News, 91, 32.[Web of Science ®] , [Google Scholar] and the Tanytarsus ortoni-group were recorded for the first time in the state of Pernambuco, while Nanocladius Kieffer, 1913a Kieffer, J.J. (1913b), ‘Nouvelle étude sur les Chironomides de l'Indian Museum de Calcutta’, Records of the Indian Museum, 9, 119–197. [Google Scholar] was recorded for the first time in the Northeast Region of Brazil. Our results make evident how much and where current knowledge of the northeastern Brazil chironomids remains fragmentary.     

The morphology of the larval head of Tipulidae (Diptera, Insecta) - The dipteran groundplan and evolutionary trends

External and internal head structures of the larva of Tipula montium are described in detail. The results are compared to conditions found in other representatives of Tipuloidea and other dipteran and antliophoran lineages. Despite of the conceivably basal position of Tipulomorpha within Diptera, the larvae are mainly characterised by derived features. The partially retracted head, the specific hemicephalic condition and several other derived character states support the monophyly of Tipuloidea. A clade comprising Tipuloidea excluding Pediciidae is suggested by the strongly retracted head, by deep dorsolateral incisions of the head capsule, by a distinctly toothed anterior premental margin, by the loss of the second extrinsic maxillary muscle, and possibly by the loss of the pharyngeal filter. Eriopterinae and Hexatominae are characterised by a tendency towards an extreme reduction of the head capsule. Limoniinae, Cylindrotomidae, and Tipulidae form a clade supported by the presence of a premaxillary suture. This implies the non-monophyly of Limoniidae. A feature shared by Cylindrotomidae and Tipulidae is the presence of a movable lacinia mobilis. However, this is arguably a plesiomorphic feature, as it also occurs in Nannochoristidae. Features of the larval head of Trichoceridae, which were included in Tipulomorpha, do not show affinities with those of Tipuloidea. Trichocerid larvae share a specialised subdivided mandible with larvae of psychodomorph groups. Tipuloidea are a highly specialised group. The characters examined did not reveal plesiomorphic features supporting a basal position, and features suggesting closer affinities with Brachycera are vague. The evolution of dipteran larval head structures was apparently strongly affected by the loss of legs and the tendency to live in cryptic habitats. Diptera are the group of Endopterygota with the highest number of apomorphic features of the larval head. The appendages are generally simplified and the muscular apparatus is strongly reduced. Specialised features evolving within dipteran lineages include specifically arranged brushes of hairs on the labrum and epipharynx, movable messores, subdivided mandibles, different mandibular brushes, and a far-reaching reduction of labial parts.     

Taxonomy and diversity of Afroalpine Chironomidae (Insecta: Diptera) on Mount Kenya and the Rwenzori Mountains, East Africa

Aim Anthropogenic climate change is expected to result in the complete loss of glaciers from the high mountains of tropical Africa, with profound impacts on the hydrology and ecology of unique tropical cold‐water lakes located downstream from them. This study examines the biodiversity of Chironomidae (Insecta: Diptera) communities in these scarce Afroalpine lake systems, in order to determine their uniqueness in relation to lowland African lakes and alpine lakes in temperate regions, and to evaluate the potential of Afroalpine Chironomidae as biological indicators to monitor future changes in the ecological integrity of their habitat. Location Mount Kenya (Kenya) and Rwenzori Mountains (Uganda). Methods The species composition of Afroalpine chironomid communities was assessed using recent larval death assemblages extracted from the surface sediments of 11 high‐mountain lakes between 2900 and 4575 m. Results were compared with similar faunal data from 68 East African lakes at low and middle elevation (750–2760 m), and with literature records of Chironomidae species distribution in sub‐Saharan Africa, the Palaearctic region and elsewhere. All recovered taxa were fully described and illustrated. Results The 11‐lake analysis yielded 1744 subfossil chironomid larvae belonging to 16 distinct taxa of full‐grown larvae, and three taxa of less differentiated juveniles. Eleven of these 16 are not known to occur in African lakes at lower elevation, and eight taxa (or 50% of total species richness) appear restricted to the specific habitat of cold lakes above 3900 m, where night‐time freezing is frequent year‐round. The faunal transition zone coincides broadly with the Ericaceous zone of terrestrial vegetation (c. 3000–4000 m). Snowline depression during the Quaternary ice ages must have facilitated dispersion of cold‐stenothermous species among the high mountains of equatorial East Africa, but less so from or to the Palaearctic region via the Ethiopian highlands. Main conclusions Chironomid communities in glacier‐fed lakes on Africa's highest mountains are highly distinct from those of lowland African lakes, and potentially unique on a continental scale. By virtue of excellent preservation and their spatial and temporal integration of local community dynamics, chironomid larval death assemblages extracted from surface sediments are powerful biological indicators for monitoring the hydrological and ecological changes associated with the current retreat and loss of Africa's glaciers.     

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 dragonflies (Odonata) in freshwater

Larvae of almost all of the 5,680 species of the insect order Odonata (dragonflies and damselflies) are dependent on freshwater habitats. Both larvae and adults are predators. The order is relatively well studied, and the actual number of species may be close to 7,000. Many species have small distributional ranges, and are habitat specialists, including inhabitants of alpine mountain bogs, seepage areas in tropical rain forests, and waterfalls. They are often successfully used as indicators for environmental health and conservation management. The highest diversity is found in flowing waters in rain forests of the tropics, the Oriental and Neotropical regions being the most speciose. This paper discusses diversity, summarises the biogeography of dragonflies in the different biogeographical regions and gives the total number of species and genera per family per biogeographical region. Examples are given of areas of particular diversity, in terms of areas of endemism, presence of ancient lineages or remarkable recent radiations but no well-based review of areas with high endemism of dragonflies is available so far. The conservation status of dragonflies is briefly discussed. Species confined to small remnants of forest in the tropics are most under threat of extinction by human activities. Guest editors: E. V. Balian, C. Lévêque, H. Segers and K. Martens Freshwater Animal Diversity Assessment     

Preliminary list of horse flies (Diptera, Tabanidae) of Serbia

Thirty six species of horse flies (Tabanidae) were previously known from Serbia (Europe). The present faunistic study of horse flies (Tabanidae) has resulted in the recording of the 4 new species Atylotus fulvus (Meigen, 1804); Tabanus miki Brauer in Brauer and Bergenstamm, 1880; Tabanus unifasciatus Loew, 1858; and Heptatoma pellucens (Fabricius, 1776), in the fauna of Serbia. The genus Heptatoma Meigen, 1803 is cited for the first time in the fauna of Serbia. 40 species are currently known from Serbia, belonging to nine genera. The fauna can be considered relatively poorly studied. Most of the species belong to the Boreal-Eurasian type of fauna 23, followed by the South European group with 8 species, the Mediterranean group with 6 species, European group with 2 species and Central European group with 1 species.