Apple snails (Ampullariidae: Pomacea) native to the New World have become agricultural and environmental pests widely in southern and eastern Asia since their introduction in about 1980. Although their impacts have been extensively documented, considerable confusion persists regarding their identities and geographical origins. Efforts to resolve the confusion have suffered from inadequate taxonomic and geographical sampling from both native and introduced ranges. Using phylogenetic and genealogical methods, we analysed 610–655 bp of cytochrome c oxidase subunit I DNA sequences from 783 apple snails from 164 Asian locations and 57 native South American locations. In Asia, we found four species of Pomacea in two clades: (1) Pomacea canaliculata and P. insularum, and (2) P. scalaris and P. diffusa. Parsimony networks and mismatch distributions indicate that the non‐native ranges of the two most widespread species, P. canaliculata and P. insularum, probably result from multiple introductions. Molecular analyses are consistent with early accounts; non‐native P. canaliculata populations trace back to multiple locations in Argentina and have probably been introduced more than once. In contrast, P. insularum was probably introduced from Brazil and Argentina independently. Multiple introductions may, in part, explain the success and rapid spread of these two species. Unlike P. canaliculata and P. insularum, P. scalaris and P. diffusa were probably introduced through the aquarium trade, derived originally from Argentina and Brazil, respectively. Possible physiological, ecological, and native range differences among these four species highlight the importance of accurate identification in understanding invasion patterns and processes, which is vital in developing and implementing management strategies.
Ampullariidae comprises two lineages of freshwater gastropods: one Old World and one New World. Recent molecular work confirmed the monophyly of the family and began to clarify generic relationships, but current systematics remains unsatisfactory. With more than 300 available species group names for New World taxa alone, taxonomic confusion is rampant, as illustrated by two species that have been introduced widely and are difficult to differentiate conchologically, Pomacea maculata Perry, 1810 and Pomacea canaliculata (Lamarck, 1822). Misidentification hampers efforts to manage their spread and impacts as invasives, and prevents meaningful comparative analyses of their biology. Here we clarify the taxonomy, describe the morphological and genetic distinctiveness of the two species, and re-evaluate their biogeographic ranges. They differ most clearly genetically, with no shared haplotypes and a mean genetic distance of 0.135 at cytochrome c oxidase subunit I (COI). Differences in shell morphology are most obvious in recently hatched juveniles; the number of eggs per clutch is higher in P. maculata, and the individual eggs are smaller, so P. canaliculata hatchlings are nearly twice as large as those of P. maculata. Adult shells differ primarily in the angulation of the whorl shoulder and pigmentation of the inner pallial lip, with the latter a distinctive feature of P. maculata. They also differ in reproductive anatomy, most notably in P. canaliculata having two distinctive glandular tissues in the apical penial sheath gland, and P. maculata lacking a medial sheath gland but possessing a basal sheath gland. Pomacea canaliculata is restricted to a narrower southern range, whereas P. maculata ranges extensively throughout much of South America. Ampullaria gigas Spix, 1827 and Ampullaria insularum d'Orbigny, 1835 are herein synonymized with P. maculata. Neotypes are designated for P. maculata and A. gigas, and a lectotype is designated for A. insularum. A neotype is designated for P. canaliculata.
Apple snails (Ampullariidae) are a diverse family of pantropical freshwater snails and an important evolutionary link to the common ancestor of the largest group of living gastropods, the Caenogastropoda. A clear understanding of relationships within the Ampullariidae, and identification of their sister taxon, is therefore important for interpreting gastropod evolution in general. Unfortunately, the overall pattern has been clouded by confused systematics within the family and equivocal results regarding the family's sister group relationships. To clarify the relationships among ampullariid genera and to evaluate the influence of including or excluding possible sister taxa, we used data from five genes, three nuclear and two mitochondrial, from representatives of all nine extant ampullariid genera, and species of Viviparidae, Cyclophoridae, and Campanilidae, to reconstruct the phylogeny of apple snails, and determine their affinities to these possible sister groups. The results obtained indicate that the Old and New World ampullariids are reciprocally monophyletic with probable Gondwanan origins. All four Old World genera, Afropomus, Saulea, Pila, and Lanistes, were recovered as monophyletic, but only Asolene, Felipponea, and Pomella were monophyletic among the five New World genera, with Marisa paraphyletic and Pomacea polyphyletic. Estimates of divergence times among New World taxa suggest that diversification began shortly after the separation of Africa and South America and has probably been influenced by hydrogeological events over the last 90 Myr. The sister group of the Ampullariidae remains unresolved, but analyses omitting certain outgroup taxa suggest the need for dense taxonomic sampling to increase phylogenetic accuracy within the ingroup. The results obtained also indicate that defining the sister group of the Ampullariidae and clarifying relationships among basal caenogastropods will require increased taxon sampling within these four families, and synthesis of both morphological and molecular data.
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