Comparing apples with apples: clarifying the identities
of two highly invasive Neotropical Ampullariidae
(Caenogastropoda)
KENNETH A. HAYES
1
*, ROBERT H. COWIE
1
, SILVANA C. THIENGO
2
and
ELLEN E. STRONG
3
1
Center for Conservation Research and Training, Pacific Biosciences Research Center, University of
Hawaii, 3050 Maile Way, Gilmore 408, Honolulu, HI 96822, USA
2
Instituto Oswaldo Cruz/Fiocruz, Av. Brasil 4365, 2104-900 Rio de Janeiro, RJ, Brasil
3
Smithsonian Institution, National Museum of Natural History, P.O. Box 37012, MRC 163,
Washington, DC, WA 20013-7012, USA
Received 3 May 2012; revised 10 August 2012; accepted for publication 22 August 2012
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.
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166, 723–753.
doi: 10.1111/j.1096-3642.2012.00867.x
ADDITIONAL KEYWORDS: anatomy – apple snails – biogeography – invasive species – lectotype –
mitochondrial DNA – Mollusca – neotype – Pomacea – taxonomy.
INTRODUCTION
The Ampullariidae (apple snails) are freshwater
gastropods found throughout the humid tropics and
subtropics. They are a key group of basal caenogas-
tropods, the largest group of gastropods. They play
important ecosystem, disease vector, and agricultural
pest roles, and have great potential as models in
evolutionary biology (Hayes et al., 2009a; Thiengo
et al., 2011). The family originated in Gondwana, in *Corresponding author. E-mail: khayes@hawaii.edu
Zoological Journal of the Linnean Society, 2012, 166, 723–753. With 14 figures
© 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 166, 723–753 723