Taxonomy of golden apple snails (Ampullariidae).

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TAXONOMY OF GOLDEN APPLE SNAILS (AMPULLARIIDAE) 1
1Department of Biology, College of Science, University of the Philippines Baguio, Baguio City; E-mail:
zbaoanan@yahoo.com
2Institute of Biology, College of Science, University of the Philippines Diliman, Quezon City; E-mail:
roberto_pagulayan@yahoo.com
Abstract
A historical account of the apple snails present in the Philippines is presented.
Although fossil data are scanty at best, they suggest that apple snails probably arose
in the Eocene period in Africa. They are now known to be pantropical in distribution.
At least 14 genera are recognized, 2 of which are native to the Philippines. The
introduced species — Pomacea canaliculata — has gained notoriety as a major pest in
rice fields, not only in the Philippines but in other Asian countries as well.
Key words: ampullariidae, golden apple snails, taxonomy
Introduction
Information on the taxonomy of the Philippine snails belonging to the family
Ampullariidae (referred to as Pilidae by some authors) is very deficient. There is a dearth of
critical scientific data available aside from the initial reports on ampullariids from the
Philippines written a century and a half ago by Philippi (1851) and Reeve (1856). Although a
number of species have been named and described, an accurate account of the number of
existing species and a more reliable system of identification have yet to be clearly established.
Species differentiation has been based primarily on a very limited set of shell samples with
rather polymorphic conchological features, resulting in a proliferation of names. A number of
Taxonomy of Golden Apple
Snails (Ampullariidae)
Zenaida G. Baoanan1
and Roberto C. Pagulayan2

2 GLOBAL ADVANCES IN ECOLOGY AND MANAGEMENT OF GOLDEN APPLE SNAILS
factors may have contributed to such confusion, mainly: (a) earlier investigators/collectors
being of different nationalities, and specimens collected being housed at different museums;
(b) the limited number of specimens collected on which descriptions were based; and (c) the
nonconsideration of the soft anatomical parts as being of taxonomic value.
Scientists are beginning to recognize these flaws and are shifting towards the incorporation
of modern methods of systematic analysis that employ genetic and molecular tools. Elucidation
of systematic relationships requires inputs ranging from classical comparative anatomy and
shell morphology to genetic and molecular data.
Family Ampullariidae
The Ampullariids, particularly Pomacea, are commonly called “golden snail,” “golden
apple snail,” “apple snail,” “jumbo snail,” “golden miracle snail,” or “giant apple snail”
(Mochida 1991). In the Philippines, they are known as kuhol or cohol in Tagalog and bisocol in
Ilocano and Pangasinan. Cowie (1995) stated that the use of the name “golden” can be
confusing, because there can be considerable color variation within a single species. Individuals
can be green, brown, orange, yellow, or many intergrades. They can also have dark brown
superimposed on the basic color. The words “giant” or “jumbo” refer to their size, as they are
the largest of ampullariid species reaching up to more than 100 mm (TROPMED Technical
Group 1986) and resembling the shape of an apple. Howells (2001) stated that name “golden”
apple snail for Pomacea is not in reference to their color but to the amount of money snail
wranglers expected to make raising them. The American Fisheries Society has recommended
the use of “channeled applesnail” (not apple snail) for P. canaliculata, with other species being
spiketop applesnail (P. bridgesii), Florida applesnail (P. paludosa), and titan applesnail (P.
haustrum) (Howells and Smith 2002).
Historical Account
Pain (1972) reported an historical survey of Ampullariids. The fossil record is scanty and
confined to areas in which the family is still found living. Earliest definite records are from
the Lutetian (Mid-Eocene) of the Faiyum of Egypt and from the Miocene of Kenya. Material
from the Cretaceous of the south of France referred to this family, but has since been shown to
belong to Natica. Only a single extinct species of the hyperdextral genus Lanistes occurred in
the Eocene of Egypt (L. antiquus Blanckenhorn 1901), probably representing the ancestral
form of L. carinatus (Olivier). Both these species are found living in the area, a tribute to the
longevity of the family in East Africa. In India, the occurrence of subfossil opercula of Pila in
Kashmir, on which Prashad (1925) based a new species of P. prisca, points to a greatly extended
range for this family within the Indian subcontinent in former times. In Africa also, Pleistocene
deposits in the Sahara containing Pila indicate a far wider range northwards than exists today,
the genus now being confined in North Africa to the Nile valley.
Distribution
Pain (1972) likewise reported the distribution of this family. Ampullariids are said to be
entirely confined to tropical areas. In America their range extends from Central Mexico to the
La Plata system, one species occurs in Florida and Georgia, and a few are found in the West
Indies. In Africa they extend along the Nile into Lower Egypt and are most abundant in the

TAXONOMY OF GOLDEN APPLE SNAILS (AMPULLARIIDAE) 3
eastern and central portions of the continent. In West Africa they do not appear to have been
found north of Lake Chad and the Niger. In South Africa the southernmost records are from
Damaraland and Bechuanaland in the west and the mouth of the Zambezi in the east. Two
species are known from Madagascar. Within the Oriental region Ampullariidae occur in Sri
Lanka, India, the basin of the Ganges, Myanmar, Thailand, Malaysia, and the islands of the
Malay Archipelago, as far east as Bali and the Celebes and northwards to the Philippines.
Some species that are indigenous in some localities were introduced to other areas for
their supposed value as food for humans and ducks, as reported by Mochida (1991). Pomacea
canaliculata (Lamarck), a species indigenous to South America, was introduced to Taiwan from
Argentina in 1979-1980 and into Kyuusyuu and Wakayama, Japan from Taiwan in 1981. In
the Philippines, P. canaliculata was introduced from Taiwan into the Rafael Atayde Hatchery,
Lemary, Batangas, Luzon in 1982. P. gigas (Spix) was imported by the Bio-Research Institute,
Metro Manila, from Florida, USA in 1983. P. cuprina (Reeve) was also introduced to Manila
around 1983. P. canaliculata was introduced from Luzon around 1983 and directly from
Argentina to Taiwan into the Asturias Farm in Cebu in 1984 for commercial production. Pomacea
culture was recommended to farmers through a livelihood project of the Philippine Government
up to 1988.
Classification of Ampullariid snails
The Ampullariidae are a group of freshwater prosobranch gastropods known to science
since pre-Linnean times (Michelson 1961). However, the classification scheme for all animals
accepted today originated in the tenth edition of Linnaeus’ Systema Naturae per Regna tria
Naturae (1758) ( Röckel et al. 1995).
Below is the outline of the classification of the ampullariids (TROPMED Technical Group,
1986):
Kingdom Animalia
Phylum Molluca
Class Gastropoda Cuvier 1797
Subclass Prosobranchia Milne Edwards 1848
Order Mesogastropoda Thiele 1927
Superfamily Viviparoidea Gray 1847
Family Ampullariidae Guilding 1828
Cowie (1997) proposed to the International Commission on Zoological Nomenclature as
Case 2996, published in the Bulletin of Zoological Nomenclature, the family name Ampullariidae
Gray 1824 as the nomenclaturally valid synonym of Pilidae Preston 1915. As a background of
this proposal, Cowie mentioned that the family Ampullariidae was introduced by Gray in
1824 (p. 276) as “Ampullariadae” based on Ampullaria Lamarck and including both Old and
New World taxa. Cowie did not mention who changed the name Ampullariadae to
Ampullariidae. The TROPMED Technical Group (1986) adopted the family name
Ampullariidae Guilding 1828 but did not write the source in their list of references. Pain
(1972) cited the work of Guilding in 1828 in his review paper entitled The Ampullariidae, an
historical survey. Pain inadvertently wrote 1802 in the text instead of 1828. Unfortunately, Pain
only mentioned Guilding’s effort to discriminate Pila and Pomacea and did not refer to any
changes in the family name. It can be deduced based from these works that it was Guilding
who first used the family name Ampullariidae as referring to the Ampullariadae by Gray
1824. Cowie (1997) further mentioned in his proposal that Preston (1915, p. 96), acknowledging
the synonymy of Pila Röding 1798 and Ampullaria Lamarck 1799, and the priority of the former,

4 GLOBAL ADVANCES IN ECOLOGY AND MANAGEMENT OF GOLDEN APPLE SNAILS
introduced the name Pilidae (Preston 1915) as a new name for Ampullariidae, perhaps, although
not explicitly, considering Ampullariidae invalid because the generic name on which it was
based was a junior synonym. Since the names of their type genera are objectively synonymous,
the same is true of the family names. Cowie argued that in Article 40b of the code it states
that, if the family-group name (in this case Ampullariidae) had been replaced before 1961
because the generic name on which it was based is recognized as a junior synonym, then the
replacement name (in this case Pilidae) is to be maintained providing that it has “won general
acceptance.” Ampullariidae is the earlier name; Pilidae has been widely used and remains in
use by some authors, possibly in the mistaken belief that usage of a family-group name is
dependent on its name-bearing genus being currently used as a valid name. Arguably, however,
the name Pilidae, although widely used, has not won “general acceptance.” Ampullariidae
also continues to be used. Lists of 48 representative publications using Ampullariidae and 21
publications using Pilidae are held by the Commission Secretariat; a single publication gave
both names as equally valid. The name Ampullariidae currently appears to be the more widely
used. In retaining the earlier name Ampullariidae, the interests of stability would be served
in a group that is likely to attract increasing attention, as some of its members (particularly the
Pomacea) are becoming serious pests of rice and other crops in Southeast Asia.
The International Commission on Zoological Nomenclature accordingly confirmed that
the family-group name Ampullariidae Gray 1824 is the nomenclaturally valid synonym of
Pilidae Preston 1915. The decision was published as Opinion 1913 in the Bulletin of Zoological
Nomenclature 56(1) March 1999. The name Ampullariidae Gray 1824, type genus Ampullaria
Lamarck, 1799 (a junior objective synonym of Pila Röding 1798), was thereby placed on the
Official List of Family-Group Names in Zoology, while the name Pilidae Preston 1915 was
thereby placed on the Official Index of Rejected and Invalid Family-Group Names in Zoology.
Snails in this family are generally large, adults being 25 mm up to more than 100 mm in
length, globose, subglobose or oval, with spire usually depressed, less than half of the total
shell length; aperture elongate, or round, with concentric calcareous operculum (TROPMED
Technical Group 1986). They are generally adapted to an amphibious mode of life by possession
of a gill and accessory lung sac inside the mantle cavity (Andrews 1965). Because of this,
some snails can be collected from land and are mistaken as terrestrial gastropods. Linné for
one referred to the common oriental species Pila ampullaceal as the genus Helix (Pain 1972).
Their possession of lungs also triggered questions whether they are primarily pulmonates
with secondary resemblance to the prosobranchs, or primarily prosobranchs with secondary
resemblance to the pulmonates (Brooks and McGlone 1908). It is not surprising, therefore,
that some authors classify them under Pulmonata (Rosario and Moquete 2002). However,
Brooks and McGlone (1908) showed that the lung of Ampullaria is a member of the series of
gill-filament, and it must be regarded as a modified filament or more than one filament. It is
thus a secondary acquisition that is not derived from the lung of the pulmonates. There is no
reason to think, therefore, that there is an ancestral connection or relationship between the
lung of Ampullaria and that of pulmonates. The structures of Ampullaria also indicate that they
are true prosobranchs.
Berthold (1991) divided the Family Ampullariidae into two subfamilies based on character
evidence. These are the monotypic Afropominae, containing just a single recent species, and
the Ampullariinae. He subdivided the Ampullariinae into the tribes Sauleini (monotypic)
and Ampullariini, with the latter further subdivided into the groups Heterostropha and
Antlipneumata.
The family Ampullariidae includes 10 genera and some 100 species according to Cowie
(1997). Although there are more than ten genera reported in the literature, some are already
invalid names. These valid and invalid names are listed in Table 1 with a discussion following.

TAXONOMY OF GOLDEN APPLE SNAILS (AMPULLARIIDAE) 5
Table 1. List of reported genera under family Ampullariidae and their respective distribution.
Genus Distribution Source
Lanistes Monfort, 1810 Africa Michelson, 1961; Berthold, 1991
Saulea Gray 1867 Africa Michelson 1961, Berthold, 1991
Afropomus Pilsbry &
Bequaert, 1927 Africa Michelson 1961; Berthold 1988, 1991
Pila Bolten 1798 Africa and Asia Prashad 1925, Michelson 1961
Pila Röding 1798 Africa Pain 1961
Pomacea Perry, 1810 South America, Central Michelson 1961
America, West Indies, Berthold 1991
southern US
Marisa Gray 1840 South America, West Indies Michelson 1961, Berthold 1991
Asolene D’Orbigny 1837 South America Michelson 1961, Berthold 1991
Pomus Sowerby 1842 China (?) Humphrey 1797 (cf Dall 1904)
Turbinicola Annandale & Asia Berthold 1991
Prashad 1921 India Prashad 1925
Pachylabra Swainson 1840 India Annandale 1920
AmpullariaLamarck 1799 Africa and Asia Dall 1904
Pomella Gray 1847 South America Berthold 1991
Felipponea Dall 1919 South America Berthold 1991
Forbesopomus Bequaert
& Clench, 1937 Philippines Bequaert & Clench 1937
Michelson (1961) attempted to divide the family Ampullariidae into two major groups, the
Old World and the New World, based on the characteristics of the kidney and penial complex.
Lanistes, Saulea, and Afropomus found in Africa and Pila from Africa and Asia are regarded as
Old World forms. Pomacea in South and Central America, the West Indies, and the southern
US, Marisa in South America and the West Indies; and Asolene, mainly in South America are
regarded as New World forms. Pila and most Old World taxa have a calcareous operculum
and a brevisiphonate siphon (i.e., the siphon is no longer than broad and distinctively funnel-
shaped). All New World genera, however, have a horny operculum, and the siphon is
longisiphonate (i.e., the siphon when expanded is an elongated cylindrical tube many times
as long as thick).
The generic name Pomus was proposed by Humprey in 1797 but was questioned by Dall
(1904) who argued that Humprey’s work was published anonymously, was not issued for sale,
bears neither the name of the author nor the publisher, and is invalid under the rules of
nomenclature. Dall further noted that, although five nude specific names were given by
Humprey (i.e., no diagnoses nor figures were cited) in his list, only one of them was identifiable,
and that was Pomus ampullaceal from China, to which is added a synonym “Helix ampullaceal.”
Dall (1904) developed a key to trace the subdivisions of the genus Ampullaria. He regarded
Saulea as a subgenus of Pila. He further subdivided the genus Ampullaria into three sections:
Ceratodes, Limnopomus, and Pomella. Under the genus Ampullaria is the subgenus Asolene, and
finally under the genus Lanistes is the subgenus Meladonus.

6 GLOBAL ADVANCES IN ECOLOGY AND MANAGEMENT OF GOLDEN APPLE SNAILS
Among the Indian ampullariids, Pila Bolten 1798 are those found in the plains while
Turbinicola Annandale and Prashad 1921 are those found in the hill-stream (Prashad 1925).
The conflict in the authorship of Pila was recalled by Cowie (1997). The Museum
Boltenianum (1798), in which the name Pila first appeared, was a catalogue compiled by J.F.
Bolten of his large mollusk collection. It was prepared for publication as a sales catalogue
after Bolten’s death by P.F. Röding, who added the synonyms used in Gmelin’s [1791] edition
of Linnaeus’ Systema Naturae and references to published figures. New generic names published
in the work were largely ignored in the 19th and first part of the 20th centuries, because very
few copies of the work were known before the publication of a facsimile edition (of 75 copies)
in 1906; the new genera were not described, their identity being determined only by the included
nominal species, which were themselves recognized only by bibliographic references to
published figures; many authors considered that new names published in sale catalogues
were not available; and adoption of the new names would have cast into synonymy many of
the familiar names introduced by Lamarck the following year. Dall (1904) was the first author
to adopt the names published in the Museum Boltenianum, but it was not until 1926 (Opinion
96) that the work was accepted by the Commission as being available. It was placed on the
Official List in 1956 (Direction 48) with authorship attributed to P.F. Röding. No diagnosis
was given for Pila, but the included species were accompanied by references to the names,
publications, and illustrations of previous authors. Pila Röding 1798 is therefore available
under Articles 12b(5) and 12b(7) of the Code.
Pila was the only genus of Family Ampullariidae represented in South and Southeast
Asia (TROPMED Technical Group 1986) until the genus Pomacea was introduced in the region.
In the Philippines, however, a new genus Forbesopomus, was reported by Bequaert and Clench
(1937). The names Pila Röding 1798 and Pomacea Perry 1810 are the names refer, respectively,
to the Old and New World genera of freshwater operculate gastropods known as apple snails.
There are confusions raised in the use of the name Ampullaria Lamarck 1799, a junior objective
synonym of Pila, because it is sometimes used for both the Old and New World species, while
Ampullarius de Montfort 1810 has also been used for New World taxa. This persistent confusion
in the literature in the generic names noted by Cowie (1997) prompted him to apply for the
placement of the names Pila Röding 1798 and Pomacea Perry 1810 on the Official List of Generic
Names in Zoology and the rejection of Ampullaria Lamarck 1799 and Ampullarius de Montfort
1810. These applications appeared together with the proposed validation of the Family
Ampullariidae. As apple snails were becoming increasingly serious pests, it was important to
establish unambiguous names for the Old and New World genera, as it was for the family
names. The case was approved, and the names Pila Röding 1798 (gender: feminine) (senior
objective synonym of Ampullaria Lamarck 1799), type species by subsequent designation by
Dall (1904) Helix ampullaceal Linnaeus 1758; and Pomacea Perry 1810 (gender: feminine), type
species by monotypy Pomacea maculata Perry 1810 were placed on the Official List of Generic
and Specific Names in Zoology. The name Ampullaria Lamarck 1799 (a junior objective synonym
of Pila Röding 1798) and Ampullarius de Montfort 1810 (an unjustified emendation of Ampullaria
Lamarck 1799 and a junior objective synonym of Pila Röding 1798) were placed on the Official
Index of Rejected and Invalid Generic Names in Zoology (Opinion 1913, March 1999).
Another case of an confusion in the specific name of an ampullariid is Ampullaria
canaliculata Lamarck 1822 (currently known as Pomacea canaliculata Family Ampullariidae).
Cowie and Kabat (2001) proposed to the Commission to conserve the well known and used
specific name of Ampullaria canaliculata Lamarck 1822 for a species of freshwater gastropod.
They argued that the name has been used as a taxon for nearly 180 years but is a junior primary
homonym of Ampullaria canaliculata Lamarck 1804 (currently known as Natica or Amauropsina

TAXONOMY OF GOLDEN APPLE SNAILS (AMPULLARIIDAE) 7
canaliculata, Family Naticdae or Ampullosporidae), the name for an Eocene marine species
from Europe. The species have not been considered congeneric since 1832.
Under the plenary power of the Commission on Zoological Nomenclature, it was ruled
that the specific name canaliculata Lamarck 1822, as published in the binomen Ampullaria
canaliculata, was not invalid by reason of being a junior primary homonym of Ampullaria
Lamarck 1804, and it was thereby placed on the Official List of Specific Names in Zoology,
and the name Amauropsina Chelot 1885 (gender: feminine), type species by original designation
Ampullaria canaliculata Lamarck 1804 was thereby placed on the Official List of Generic Names
in Zoology. The name canalifera Lamarck 1822, as published in the binomen Ampullaria
canalifera, was thereby placed on the Official Index of Rejected and Invalid Specific Names in
Zoology (a junior objective synonym of Ampullaria canaliculata Lamarck 1804) (Opinion 1997,
June 2002).
The taxonomic key of ampullariids has been based mainly on shell morphology (Burch
1980, Upatham et al. 1983). However, identification at the species level based solely on shell
morphology proved to be confusing, resulting in a proliferation of names. Hence, other
characters are considered for taxonomic classification. Snails are classified by using any or a
combination of the following characters:
• Shell form or conchological characteristics. The different genera of ampullariids have
been erected primarily upon conchological characteristics based on such criteria as
color, size and shape of the shell, types of sculpturing (if present), the presence or
absence of an umbilicus, and the description of the operculum (Michelson 1961).
However, this is not a solid basis, since there could be morphological variations within
the same species.
• Radula. All members of the family Ampullariidae possess a taenioglossate radula
used for feeding (i.e., with two marginal and one lateral teeth on both sides of the
central tooth, giving the radular formula 2.1.1.1.2). There seem to be no significant
differences between the radula of the different ampullariids so far studied; hence it
can be of taxonomic value only in the identification of the family but not of the genera
nor the species (Demian 1964, Pagulayan 1991).
• Soft parts. The study of soft parts or anatomy of snails is done to elucidate intra- and
interspecific variation in different species. Berthold (1991) assessed the comparative
analysis of 36 species from 6 genera of the Ampullariidae. His results, supplemented
by data from the literature, allowed him to reconstruct ampullariid phylogeny.
• Cytogenetics
Genus Pomacea
Of all the ampullariids, Pomacea has become the most important, as it is becoming a
major pest throughout most of South and Central America, the Caribbean, and Southeast
Asia. There are about 50 species of Pomacea (Cowie, in press) with the possibility of homonymy.
Some of these species available in the literature are listed in Table 2. Future taxonomic revision
may well synonymize species within this group.

8 GLOBAL ADVANCES IN ECOLOGY AND MANAGEMENT OF GOLDEN APPLE SNAILS
Table 2. Pomacea species and their possible origin. The species with
no reference were cited by Cowie (in press) in his review paper.
Species Origin Reference
Amazonica South America
australis ? Madamba and Camaya 1987
bridgesii South America Cowie 1993, Howells 2001
canaliculata Argentina Scott 1957, Berthold 1991
columellaris Peru Michelson 1961
cuprina Mochida 1987
decussata
dolioides Surinam van Dinther 1973
falconensis
fasciata
flagellate Central America
gigas Mochida 1987
glauca Guadalupe Berthold 1991
gossei
guyanensis
hanleyi
haustrum Brazil Guimarães 1983
immersa
insularum Argentina Scott 1957
intermedia Brazil Berthold 1991
interrupta Peru Michelson 1961
lineata Brazil Thiengo 1987
luteostoma Venezuela (?)
megastoma
nais
nublia Peru Michelson 1961
paludosa Florida, US Hanning and Leedon 1978
papyracea South America
pyrum
scalaris Argentina Scott 1957
sordida Brazil Thiengo 1989
urcea Brazil Berthold 1991
urceus Venezuela Burky 1974

TAXONOMY OF GOLDEN APPLE SNAILS (AMPULLARIIDAE) 9
Golden Apple Snails Reported in the Philippines
There remains some confusion over the identification and nomenclature of the so-called
golden apple snails reported in the Philippines. Various names reported include Ampullaria
gigas, Pomacea sp, Pila leopordvillensis (Guerrero 1991); Ampularius sp. (Lacanilao 1990); Pomacea
insularis (Acosta and Pullin 1991); P. gigas, P. cuprina, and P. canaliculata (Mochida 1991).
Specimens collected from rice fields at the Central Luzon State University and sent to the
Department of Malacology, Academy of Natural Sciences in Philadelphia, US were identified
by Dr. George Davis as P. insularis. The Food and Agriculture Commission and the
Commonwealth Agricultural Bureau-International Institute of Biological Control likewise
identified them as P. insularis. However, based on the morphological characteristics, they were
identified as P. canaliculata at the International Rice Research Institute in Los Baños, Laguna
(Saxena et al. 1987). Most of the published reports refer to golden apple snails in the Philippines
as P. canaliculata (Cagauan and Joshi 2002).
The diagnostic characteristics that will distinguish between the introduced Pomacea and
the native species of Pila are summarized in Table 3.
Table 3. Diagnostic characteristics between Pomacea and Pila.
Diagnostic Pomacea Pila Reference
Feature
Shape of shell dextral; sub-ovate, dextral; sub-ovate Michelson 1961
ovate, or globose to globose
Umbilicus open umbilicus closed to slightly Pagulayan 1991
open
Operculum corneus or horn-like calcareous, hard, Michelson 1961
and pliable and brittle
Respiratory siphon longisiphonate brevisiphonate Michelson 1961
Sperm canal internal external Michelson 1961
Reproductive organs pallial vas deferens glandular walls of Pagulayan 1991
continuous with the pallial vas deferens
the penis in males; form prominent ridges
proximal portion of used for species;
capsule gland is identification; vas deferens
coiled and embedded terminates into the genital
in the tissue of the papilla; bursa copulatrix
albumen gland in are found outside the
females albumen gland
Nature of eggs calcareous shell; calcareous shell; Michelson 1961
pigmented (mostly nonpigmented
pink to red) (white)
Oviposition site on emergent parts in banks or mudflats Michelson1961
of aquatic vegetation
Behavior moderately highly Michelson 1961
amphibious amphibious

10 GLOBAL ADVANCES IN ECOLOGY AND MANAGEMENT OF GOLDEN APPLE SNAILS
Conclusion
The taxonomy of golden apple snails in the Philippines still warrants special attention
from the scientific community. Not much detailed taxonomic research has been done to resolve
the confusion in identifying these snails. As they continue to threaten rice production, there
must be a solid and logical way to resolve their identity before detailed ecological and
management strategies can be devised for these pests.
Acknowledgement
The authors would like to thank Dr. Ravi Joshi for providing a copy of the Scientific
Information Database on Golden Apple Snail (Pomacea spp), from which some of the reference
materials were taken.
References
Acosta, B.O., and R.S.V. Pullin. 1991. Environmental impact of the golden snail (Pomacea sp.)
on rice farming systems in the Philippines. Muñoz, Nueva Ecija: Freshwater Aquaculture
Center, Central Luzon State University, and Manila: ICLARM. vi + 34 pp.
Andrews, E.B. 1965. The functional anatomy of the mantle cavity, kidney, & blood system of
some pilid gastropods (Prosobranchia). J. Zool. 146:70-94
Annandale, D. 1920. The apple-snails of Siam. Journal of the Natural History Society of
Siam. 4(1). 19 pp with 2 plates.
Bequaert, J.C. and W.J. Clench. 1937. Forbesopomus, a new genus in the family Pilidae
(Ampullariidae), from the Philippine Islands. Proc. New England. Zool. Club 16:
53-56
Berthold, T. 1988. Anatomy of Afropomus balanoideus (Mollusca, Gastropoda, Ampullariidae)
and its implications for phylogeny and ecology. Zoomorphology. 108:149-159.
Berthold, T. 1991. Vergleichende Anatomie, Phylogenie und Historiche Biogeographie
der Ampullariidae (Mollusca, Gastropoda) (Comparative anatomy, phylogeny,
and historical biogeography of the Ampullariidae (Mollusca, Gastropoda).
Abhandlungen des Naturwissenschaftlichen Vereins in Hamburg (NF). 256 pp.
Brooks, W.K., and B. McGlone. 1908. The origin of the lung of Ampullaria. Carnegie Inst.
Publ. 102:97-111.
Burch, J.B. 1980. A guide to freshwater snails of the Philippines. Malacological Review
13(1/2):121-143.
Burky, A.J. 1974. Growth and biomass production of an amphibious snail, Pomace urceus
(Müller), from the Venezuelan savannah. Proc. Malac. Soc. Lond. 41:127-143.
Cagauan, A.G. and R.C. Joshi. 2002. Golden apple snail spp. in the Philippines. In: Wada, T.,
Y. Yusa, and R.C. Joshi. 2002. Proceedings of the Special Working Group on the Golden
Apple Snail (Pomacea spp.) Los Baños, Laguna, Philippines.
Cowie, R.H. 1993. Identity, distribution and impacts of introduced Ampullariidae and
Viviparidae in the Hawaiian Islands. J. Med. and Appl. Malacol. 5: 61-67
__________________. 1995. Report on a visit to Cambodia to advise on apple snails as potential
rice pests. In: Wada, T., Y. Yusa, and R.C. Joshi. 2002. Proceedings of the Special Working
Group on the Golden Apple Snail (Pomacea spp.) Los Baños, Laguna, Philippines.

TAXONOMY OF GOLDEN APPLE SNAILS (AMPULLARIIDAE) 11
__________________. 1997. Case 2996: Pila Röding, 1798 and Pomacea Perry 1810 (Mollusca,
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