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Global Diversity of Gastropods (Gastropoda; Mouusca) in Freshwater

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Global Diversity of Gastropods (Gastropoda; Mouusca) in Freshwater Hydrobiologia (2008) 595:149-166 DOI 10.1007/S10750-007-9012-6 'RESHWATER ANIMAL DIVERSITY ASSESSMENT Global diversity of gastropods (Gastropoda; MoUusca) in freshwater Ellen E. Strong • Olivier Gargominy • Winston F. Ponder • Philippe Bouchet ' Springer Science+Business Media B.V. 2007 Abstract The world's gastropod fauna from conti- gastropods. Despite their ecological importance in nental waters comprises ~ 4,000 valid described many aquatic ecosystems, understanding of even their species and a minimum of 33-38 independent lineages systematics is discouragingly incomplete. The world's of Recent Neritimorpha, Caenogastropoda and Het- freshwater gastropod fauna faces unprecedented erobranchia (including the Pulmonata). The caeno- threats from habitat loss and degradation and intro- gastropod component dominates in terms of species duced fishes and other pests. Unsustainable use of richness and diversity of morphology, physiology, life ground water, landscape modification and stock dam- and reproductive modes and has produced several age are destroying many streams and springs in rural/ highly speciose endemic radiations. Ancient oligo- pastoral areas, and pose the most significant threats to trophic lakes (e.g., Baikal, Ohrid, Tanganyika) are key the large diversity of narrow range endemics in springs hotspots of gastropod diversity; also noteworthy are a and ground water. Despite comprising only ~5% of number of lower river basins (e.g., Congo, Mekong, the world's gastropod fauna, freshwater gastropods Mobile Bay). But unlike many other invertebrates, account for ~ 20% of recorded mollusc extinctions. small streams, springs and groundwater systems have However, the status of the great majority of taxa is produced the most speciose associations of freshwater unknown, a situation that is exacerbated by a lack of experts and critical baseline data relating to distribu- tion, abundance, basic life history, physiology, mor- Guest editors: E. V. Balian, C. Leveque, H. Segers and phology and diet. Thus, the already considerable K. Martens magnitude of extinction and high levels of threat Frestiwater Animal Diversity Assessment indicated by the lUCN Red List of Threatened Species is certainly a significant underestimate. E. E. Strong (El) Department of Invertebrate Zoology, Smitlisonian Institution, National Museum of Natural History, MRC Keywords Phylogeny • Taxonomy • Biogeography • 163, P.O. Box 37012, Washington, DC 20013-7012, USA Endemicity • Radiations • Life history • Fossil record • e-mail: [email protected] Biomonitoring • Disease transmission • Conservation O. Gargominy • P. Bouchet Museum National d'Histoire Naturelle, 55 Rue Buff on, 75005 Paris, France Introduction W. F. Ponder Malacology Section, Aquatic Zoology, Australian The Mollusca is an extraordinarily varied phylum— Museum, 6 College Street, Sydney, NSW 2010, Australia with estimates of 80,000-100,000 described species •^ Springer 150 Hydrobiologia (2008) 595:149-166 and total diversity possibly as high as 200,000, they and Neritimorpha (previously a subgroup of "archae- are second only to arthropods in species richness. The ogastropods"). largest molluscan classes—Gastropoda and Bival- The world's freshwater gastropod fauna is domi- via—have repeatedly and successfully colonized nated by two main components: the Caenogastropoda continental ("fresh") waters. Freshwater gastropods and pulmonate heterobranchs. Several additional are found on every continent except Antarctica and in basal lineages of heterobranchs have also invaded nearly all aquatic habitats including rivers, lakes, freshwater (Valvatidae, Glacidorbidae, Acochlidiida) streams, swamps, underground aquifers and springs, as well as some Neritimorpha (Neritiliidae, Neriti- as well as temporary ponds, drainage ditches and dae). Only the Viviparoidea, Glacidorboidea and other ephemeral and seasonal waters. Most live nearly all Hygrophila comprise superfamilial (or submerged, and many are specialized for particular above) groupings with members represented exclu- habitats—aquatic vegetation, stones, rocks, wood and sively in freshwater. Of the 409 families of Recent other solid surfaces, or soft sediment. Some are gastropods currently recognized (Bouchet & Rocroi, amphibious and a few are able to tolerate periods of 2005), 26 are composed of taxa that are wholly or time out of water (e.g., some Ampullariidae); others mostly restricted to freshwater, four have significant are capable of prolonged periods of aestivation in soil taxonomic representation in freshwater biotopes during dry periods. Few groups (notably some of the (Neritidae, Assimineidae, Hydrobiidae, Stenothyri- rissooidean families) are found in highly saline inland dae), and three are marine groups with isolated habitats such as the Caspian Sea or salt lakes in genera that have invaded freshwater [Cremnoconchus Central Asia, Africa and Australia. (Littorinidae), Clea (Buccinidae), Rivomarginella Most freshwater gastropods are micro-herbivorous (Marginellidae)] (Table 1). and/or micro-omnivorous grazers feeding on bacterial The caenogastropod component of the freshwater films, algae and diatoms, but there are a number of fauna represents numerous independent lineages and exceptions: the predominantly marine Buccinidae, many separate colonization events. Several clades Marginellidae and Acochlidiida and the entirely fresh- have produced spectacular endemic radiations, water Glacidorbidae are predators; Viviparidae and namely Rissooidea (Hydrobiidae s.L, Pomatiopsidae) Bithyniidae are ctenidial suspension feeders at least in and Cerithioidea (Pachychilidae, Paludomidae and part; Ampullariidae are primarily macroherbivorous Pleuroceridae). With the exception of a few parthe- and are also known to feed on bryozoans and planorbid nogenetic taxa [Campeloma (Viviparidae), Melano- eggs. There are no pelagic/nektonic or parasitic ides (Thiaridae), Potamopyrgus antipodarum species, with the great majority being benthic crawlers. (Hydrobiidae)], they are exclusively dioecious and A rare exception is the Helicostoidae—a monotypic reproduction is sexual. Of all the freshwater groups, caenogastropod family of uncertain affinity from China only the cerithioids are aphallate and transfer sperm that lives cemented to limestone blocks (Lamy, 1926). using spermatophores; all others use a penis. Most lay egg capsules, and development is intracapsular with Taxonomic composition embryos emerging as crawling juveniles. A free- swimming dispersal stage is present in some species, New suites of anatomical, ultrastructural and molec- particularly those that inhabit the lower reaches of ular characters developed in the past 30 years have coastal streams, with a free-swimming veliger larva fuelled a revolution in our understanding of gastropod that may develop in the sea (Neritidae, some phylogenetics (Haszprunar, 1988; Ponder & Lind- Thiaridae). However, many species are brooders [all berg, 1997; Colgan et al., 2003; Strong, 2003). Viviparidae, some Cerithioidea, Rissooidea and Het- Several well supported clades are currently recog- erobranchia (see below)] and retain their young in nized: Caenogastropoda (containing most of the brood pouches that represent modifications of the former Mesogastropoda and all the Neogastropoda); oviduct, mantle cavity or cephalic haemocoel. While its sister group, Heterobranchia (containing the it has been suggested that there is a significant former Opisthobranchia and Pulmonata, as well as a selective advantage for parental care and hence few "mesogastropod" groups); Vetigastropoda brooding among freshwater molluscs (e.g., Kohler (including many of the former Archaeogastropoda) et al., 2004), the great majority of freshwater ^ Springer Hydrobiologia (2008) 595:149-166 151 Table 1 Taxonomic representation and distribution of fresliwater gastropods Taxon Representation Habitat In Freshwater Neritimorpha Superfamily Helicinoidea Family Neritiliidae Anchialine and coastal running waters Superfamily Neritoidea Family Neritidae Primarily lower reaches of coastal rivers and streams, estuaries Caenogastropoda Architaenioglossa Superfamily AmpuUarioidea Family AmpuUariidae Quiet, muddy rivers, lakes, ponds, canals, rice paddies, swamps Superfamily Viviparoidea Family Viviparidae Rivers, lakes, ponds, swamps, canals Sorbeoconcha Superfamily Cerithioidea Family Melanopsidae Springs, streams Family Paludomidae Lakes, rivers, streams (including radiation in Lake Tanganyika) Family Pachychilidae Lakes, rivers, streams (including radiation in Sulawesi lakes) Family Pleuroceridae Rivers, streams Family Thiaridae Rivers, streams Hypsogastropoda Superfamily Littorinoidea Family Littorinidae (Cremnoconchus) Waterfalls Superfamily Rissooidea Family Amnicolidae Rivers and streams Family Assimineidae Estuaries, freshwater rivers and streams, springs Family Bithyniidae Quiet muddy rivers, lakes, ponds, canals, swamps Family Cochliopidae Rivers and streams, swamps, lakes Family Helicostoidae (Helicostoa) Cemented on limestone rocks Family Hydrobiidae Greatest diversity springs; also streams and rivers, lakes, groundwater systems, caves, estuarine marshes and mudflats Family Lithoglyphidae Streams, rivers Family Moitessieriidae Groundwater systems, caves Family Pomatiopsidae */< Rivers, permanent wetlands, stream edges, some saline springs/ lakes. Family Stenothyridae Rivers, streams, estuarine Neogastropoda Superfamily Buccinoidea Family Buccinidae (Clea) Lower reaches of rivers Superfamily Muricoidea Family Margme\[idae(Rivomarginella) Rivers, lakes
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