Freshwater Molluscan Shells

Gastropoda (Snails)
Neritinidae: Thick walled colorful shells with shelly operculum, mostly tropical, related to the marine nerites .
Ampullariidae: Large "apple snails" of the tropics world wide, generally smooth.
Viviparidae: "mystery snails", usually with circular or oval aperture, generally smooth.
Valvatidae: Small white snails, widely umbilicate.
Other freshwater snails: A few species from generally marine families
Pleuroceridae and other Cerithioidian families: Thick-walled shells, often high-spired, may have
sculpture and be colored, or with color bands. North America, Africa, Asia, South America
Thiaridae: Similar to the Pleuroceridae, mainly tropical.
Melanopsidae: Similar to the Pleuroceridae; Mediterranean region, New Zealand.
Rissoacea: A great many species of mostly small to tiny snails, world wide. Diverse in Australia.
Freshwater pulmonate snails: Thin walled shells, world-wide, usually weakly colored, may be
dextral, (Lymnaeidae)
sinistral, (Physidae), or
planispiral (Planorbidae), or restricted to certain geographic locations such as
temperate South America (Chilinidae)
or New zealand (Latiidea)

Freshwater limpets: Thin, small, saucer or cap-shaped shells, not closely related to marine limpets.

Gastropod family tree showing relationships between the major freshwater lineages.
This is a more traditional approach to classification. Recent work suggests, for instance, that the Archaeogastropoda
is an artificial grouping, the Neritinidae belong in their own order and superorder, and also that the Viviparidae and
Ampulariidae are separate lineages, both of which should connect at the base of the Caenogastropod branch.
Furthermore, Valvatoidea do not belong with the Caenogastropoda, but with Heterostropha, an order not formerly
recognized and not shown here. More recently, groups within the family Pleuroceridae have been raised to family
status, so the Cerithiacea (Cerithioidea or Vermetoidea ) now contains several additional freshwater families. Click
for
New classification in tabulated form.

Freshwater Molluscan Shells

Neritidae (Neritinidae)

Of this family of mostly tropical shallow-water marine snails, over 175 species live in fresh or
brackish waters. All graze upon algae growing on firm underwater surfaces. They are notable in
having thick variably patterned shells and a calcareous operculum. Unlike most freshwater snails
where young are born live or hatch directly from eggs, development in the nertiidae involves a

planktonic larval form referred to as a veliger. Tom Eichhorst indicates some species cannot
reproduce in freshwater aquaria, as their veligers need exposure to saltwater to develop.
Genera of the neritidae with representatives in fresh waters include:
Neritina Indo-Pacific.
Nereina Central and South America,
Theodoxus; Europe, northern Africa, Middle East.
Clypeolum; Central and South America, sub-Saharan Africa.
Septaria; Rivers and streams of Indo-Pacific islands, Ganges and Hooghly Rivers, India.
Fluvinerita; Mountain brooks of Jamaica.
Neritodryas; Indo-Pacific.
Neritilia; Circum-tropical.
Vitta; the Americas and Africa
Vittina: Indo-Pacific, southern Asia.
The ranges given above are from Banarescu, 1990, Preston, 1915, and Tom Eichhorst, who also
provided images, information, and identifications. Names and classifications are currently [2004]
under revision. Please note, some of the image filenames, where they contradict the captions,
reflect earlier classifications, or misidentification of specimens.

Western Hemisphere

"Olive Nerite"
The name Neritina reclivata (Say,
1822)
has long been used for this species,
but
Vitta usnea Rding (1798) has
priority.
Freshwater creeks in Florida, United States.
x2. Detail of banding to right, x8.

Clypeolum latissimus
(Orbigny, 1840) Rivers,
Pacific side, Equador

Clypoleum (formerly Neritina) punctulata (Lamarck, 1816). Freshwater rivers well above
marine influence, island of Dominica.
Photos, Dr. Donald J. Stewart.

Vitta (formerly Neritina)

virginea (Linne, 1758).
Vitta meleagris (Lamarck, 1822) Rio de Janiero State, Brazil

West indies.
Great variety of colors and patterns.
Photo Stefan Chessa / Germany.
Click here for photo collage.

Vitta zebra (Bruguiere, 1792) Near Belem, Brazil.

Europe
Some Theodoxux are noteworthy in
having ranges far outside of the tropics,
including England and Europe along

Other species listed for the territory of Europe

the Baltic Sea.

include:
T. anatolicus Rcluz, 1841, Turkey; Greece
T. baeticus Lamarck, 1822, Spain
T. euxinus Clessin, S., 1885, Europe
T. fluviatilis (Linnaeus, 1758), Europe
T. meridionalis Philippi, 1836, Italy
T. prevostianus Pfeiffer, 1828, Europe
T. tranversalis Pfeiffer, 1828, Europe
T. valentinus Graells, 1846, Spain
T. velascoi (Graells, 1846), Spain.

Theodoxus danubialis (Pfeiffer,1828)

Italy.
Theodoxus danubialis has acquired the
following common names:
Czech; zubovec dunajsk
German; Donau-Kahnschnecke
Hungarian; rajzos bdncsiga.

A few of the common names applied

to the wide-ranging T. fluviatilis:
Czech; zubovec rcn.
English; River Nerite.
German; Gemeine Kahnschnecke,
Swedish; Btsncka, Algsncka,
Russian; y p

Africa / Asia / South Pacific

Vittina natalensis (Reeve, 1845) African rivers, in natural state and cleaned. Image; Tom Eichhorst.

Vittina coromandeliana (G.B. Sowerby I, 1836)

left (dorsal view), and Neritodryas dubia

(Gmelin,
1791) right (aperture view). African rivers. Specimens
courtesy of Gerald Depaus, Belgium. ID; Tom Eichhorst.

Clithon cf. sowerbiana (Recluz, 1841). Okinawa, Japan.

Clithon castanea (Hombron & Jaquinot, 1854). Samoa. Image; Tom Eichhorst.

Vittina variegata (Lesson, 1831)

New Caledonia

Clithon dispar (Pease, 1867) considered to

be a subspecies of Clithon chlorostoma
(Broderip, 1832) by some authors.
Western Samoa.

Clithon corona (Linnaeus, 1758) Philippines. Markings on large

individual are concealed by algae.

Species in the fhe freshwater genus Septaria possess an internal operculum, completely buried
within the foot of the animal. Tom Eichhorst indicates that it may have the function of helping to
leverage the animal to the substrate, but it has never been shown for certain what if any function
it serves. He provides the following information, along with the images below. The operculum in
Septaria varies in size from Septaria cumingiana whose operculum is almost the same size as the
snail's aperture, to Septaria tesselata in which it appears small, thin, and vestigial. Its dark
corneous region fits behind the septum. At least four of the thirteen currently recognized
Septaria species are sexually dimorphic; S. cumingiana (Rcluz, 1842), S. porcellana
(Linnaeus, 1758), S. sanguisuga (Reeve, 1856), and S. suffreni (Rcluz, 1841). These limpet
or slipper-shell like nerites are typically found in fast flowing freshwater streams. Five species
are listed for the Indian subcontinent by Preston, 1915.

Septaria porcellana (Linnaeus, 1758), with, and without periostracum. Philippines, actual size, 26 mm.
Opercula; left, "outward" face, right, "inward" face. Photo Tom Eichhorst

Septaria suffreni (Rcluz, 1841), Samoa. With, and without periostracum. Females, top; males, bottom.
Actual size, 18 to 24 mm.
Opercula; left, "outward" face, right, "inward" face. Photo Tom Eichhorst.

Freshwater Molluscan Shells / Ampullariidae

Commonly seen in aquaria, the Ampullariidae are the large apple snails of the tropics world
wide. Please visit the apple snail site (http://www.applesnail.net) for extensive information on this
family.

Pomacea; widespread in South America, Florida

Marisa; widespread in South America
Asolene; northern South America and Parana River basin
Felipponea; lower Parana River
Pila; widespread in Africa, India, Southeast Asia
Lanistes; widespread in Africa
Saulea; west Africa
Afropomus; west Africa
Forbesopomus; Lake Lanao, Philippines

Asia

Pila ampullacea
(Linnaeus, 1758),

Pila polita
(Deshayes, 1830),

Bali.

Thailand.

Africa
Pilsbry and Bequaert (1927) in their review of the African freshwater fauna include
Pila: 21 species + 2 additional subspecies
Lanistes Subgenus Lanistes: 23 species + 3 additional subspecies
Lanistes Subgenus Meladomus: 20 species + 9 additional subspecies and varieties
Saulea: 1 species
Afropomus: 1 species.
Some of their illustrations are included below, courtesy American Museum of Natural History.

Pila ovata (G. Oliver,

1804)

Pila wernei (Philippi, 1851)

Cental Africa rivers, under water-lily pads.

central Africa.

Lanistes varicus Muller, 1774. Southwest Africa. Lanistes is not

truly sinistral, but instead, "hyperstrophic." It might be described as a
dextral shell that spirals upward as it grows, instead of downward,
such that the spire has become the umbilicus, and vice-versa.

Pila leopoldvillensis (Putzeys,

1898) P. congoensis, Pilsbry and
Bequaert (1927) is similar, with
narrower aperture. To 100 mm.

Lanistes ovum
Peters,
1845, Central
African
lakes.

Lanistes bicarinatus, Germain,

1907
Leopoldville, 39 mm.

Lanistes intortus (Lamarck

1822)
30 mm, from Zambi, Malela,
and Banana.

Lanistes procerus langi Pilsbry

and
Bequaert 1927

Salea vitrea
(Born, 1780)

Western Hemisphere

Afropomus balanoideus
(Gould, 1850)

Asolene scalaris
Lamarck, Uruguay.

Asolene megastoma
neritoidea Sowerby, 1825

Pomacea lineata
(Spix, 1827), Brazil.

Uruguay. Mature shell.

Pomacea canaliculata
(Lamarck, 1819), SE.

Pomacea paludosa
(Say, 1829), Florida, USA.

South America, common in

aquariums, invasive in SE.
Asia, Japan, Australia, Hawaii.

Pomacea insularum
d'Orbigny, 1835, Uruguay.

Felipponea neritiformis
Dall, 1919, Brazil.

Pomacea papyraceus
(Spix, 1827), South America.

Pomacea bridgesii (Reeve, 1856). Large mature shell. Amazon Basin.

Farmed locally and eaten as "churo". Small banded individuals common in aquaria.

Marisa cornuarietis (Linnaeus 1758), Florida, USA,

native to South America, also introduced into Africa.

Freshwater Molluscan Shells / Viviparidae

Viviparus georgianus (Lea, 1834), (left), and Campeloma limum (Anthony, 1860), (right). Campeloma has the large
foot characteristic of active snails that plow through sand just beneath the surface. Both species are native to
rivers and streams of the Southeastern United States coastal plain.

Viviparidae, commonly referred to as "mystery snails" have a worldwide distribution. Burch,

(1982) list 17 total species for North America. Preston, (1915) lists fourteen for India, Liu Yueyin
(1979) includes twelve for China, and Zhadin, (1952) lists five for the territory of the former
USSR. They are also present in Australia, Africa and Europe (six species, see Fischer), but
except as fossils, absent in South America. The viviparidae are noteworthy in their wide selection
of foodstuffs. Brown (1991) indicates they can function both as grazers, consuming algae
growing on any submerged surface, and detritivores, utilising fine particulate organic matter and
the bacteria and other microorginisms therein. They also filter feed on suspended matter,
competing with the clams and mussels. Dillon (2000) indicates that North American Campeloma
can also be baited with carrion, and describes the process by which Viviparus snails consume
suspended particulate material.
The gills of Viviparus are characterized by unusually large triangular lamellae whose tips hang over a ciliated gutter
or "food groove" running across the floor of the mantle cavity.... Cilia ...direct mucus and entrapped particles to the
food groove. Particles collected on the gill filaments are carried to the tip, where they also fall into the grove. A
food/mucus string forms which is carried forward and collected into a ball or "sausage". Periodically the snail will
turn its head and eat the collected food.

Principal genera, in three subfamilies:

Viviparinae;
Viviparus; Eastern North America and most of Europe.
Tulatoma; endemic in Coosa River, Alabama, USA.

Campelominae or Lioplacinae;
Campeloma; Eastern North America.
Lioplax; Eastern North America.
Bellamyinae;
Bellamya; Africa, India, southeast Asia.
Cipangopaludina; east and southeast Asia, and as an exotic elsewhere.
Notopala; eastern Australia.
Larina; northeast Australia.
unclassified;
Margarya; China
Angulyagra; China
Rivularia; China
Neothauma; Africa
Heterogen; Japan
Taia; Southeast Asia

Asia
<- Cipangopaludina
chinensis
(Reeve, 1863), now found

Amuropaludia
pachya
Bourguignat,1860,
Heterogen
longispira
Smith, Japan.

far east Russia.

near
Washington, DC, Southern
California
lakes, and other places in the
United
States and Canada. The
Chinese
mystery snail is sold in
traditional
Chinese markets.

Cipangopaludina
japonica
(Martens, 1861), also
found in the
United States, is similar, but
somewhat more high-spired
and
angular.

Thick shelled Chinese Rivularia snails: Rivularia auriculata von Martens, 1875
from Fujian Province, (above) and Rivularia bicarinata Kobelt, 1909 from Hunan
Province. (below).

Images below from Liu Yueyin (1979)

Cipangopaludina
ussuriensis
(Heude), northern
China.

Margarya
melanoides
Nevill, Yun Nan region
lakes, China.

Margarya
mansuyi
Deutzenberg et
Fischer, Yun Nan,
China.

Bellamaya
purificata (Heude),
widespread, China.

probably Angulyagra
polyzonata (Frauenfeld)
Canton, China

Southeast Asia

Filopaludina sumatrensis polygramma (Martens, 1860),

Thailand

Bellamya angularis
(Muller, 1774), Philippines

Taia elitoralis
Annadale, 1918 (left)
Anulotaia mekongensis
Brandt, 1970 (center)
Trochotaia trochoides
(Martens, 1860) (right)
Black and white images
from Brandt (1974)

Brandt (1974) lists several genera endemic to Thailand and Southeast Asia, and includes the
following numbers of species for the territory of Thailand: Filopaludina 6 species plus 4
additional
subspecies; Sinotaia 2; Annulotaia 2; Trochotaia 1; Eyriesia 1; Idopoma 3 Cipangopaludina
1; and Mekongia 6.

Europe
Please visit Wolfgang Fischer's page at
http://ipp.boku.ac.at/private/wf/Europaeische_Viviparidae_main.html for images of the six recent
and additional fossil European species.
Eng; Mystery snails
De; Sumpfdeckelschnecken
It; Viviparo, Paludina
Fr; Paludine

Ne; moerasslaken
Ru; #150;#128;
[zhivarodki]
Pol; ywordka

Viviparus
mammilatus
(?) Kuster 1852,
Monte Negro

Viviparus
viviparus
(Linnaeus, 1758)
Belgium, elsewhere in
Europe

Viviparus acerosus

(Bourguignat
1862)
Hungary

Viviparus contectus
Millet
ponds, Hungary

Africa
Pilsbry and Bequaert, (1927) list 35 Viviparus species, with 22 additional varieties, and one
species with three additional varieties for Neothauma. At least some of the Viviparus species
have since been placed with Belamya.

Neothauma tanganyicensis Bourguignat,

1885. Lake Tanganyika.
Specimen contributed by Gerald Depaus, Belgium.

Viviparus crawshayi E. A. Smith, 1893

Zaire, Africa

Viviparus mweruensis E. A. Smith, 1893,

Lake Moero, Central Africa. Shell has growth line
features in common with Neothauma tanganyicensis,
above, and "Neothauma differs primarily from
Viviparus in th shape of the outer lip" (Pilsbry and
Bequaert, 1927).

Bellamya jeffreysii
Bellamya unicolor
Frauenfeld, Central Africa Livier, 1806. Senegal

Australia

Larina strangei Adams 1854. Queensland, Australia.

Hugh Jones of the New South Wales Department of Land & Water Conservation indicates that
some of the Viviparidae are endangered in Australia. In particular, Notopala hanleyi and
Notopala sublineata from the Murray-Darling drainage. These were thought to be extinct, but a
few remnant populations have been rediscovered in irrigation pipelines. They were apparently a
problem and the last known population of N. hanleyi may have been eradicated by efforts to clear
the pipeline.

North America

Relatively high-spired Campeloma from the

Cache River in Arkansas.

Campeloma limum
(Anthony,
1860), creeks and rivers of
the
southern Atlantic coastal
plain. x 2.

Campeloma decisum (Say,

1817) Eastern United states and
southern Canada generally;
rivers and lakes.

Lioplax sulculosa (Menke, 1828)

American
mid south, larger rivers and reservoirs.

Viviparus georgianus
(Lea, 1834), southeastern
North America, apparently
invasivein Lake Champlain,
New Your.

Viviparus subpurpureus
(Say, 1829), southeastern

Viviparus intertextus
(Say, 1829), southeastern

North Anerica.

North America

Viviparid opercula are concentrially structured

and assymetrical. Shown are Viviparus
subpurpureus (left) and V. intertextus (right).

Freshwater Molluscan Shells

Hydrocenidae, Valvatidae, and other freshwater snails

Hydrocenidae
Minute amphibious snails classified with the old archeogastropoda, native to eastern and
southern Asia, and southern Europe. One species is listed for southern Africa by Pilsbry and
Becquaert (1927) "[They] are more or less amphibious dwellers in wet vegetation, clinging to the
faces of rocks continually washed by the spray from waterfalls" (Preston, 1915). He lists eight
species for the Indian subcontinent

Georissa japonica Pilsbry, 1900. x8. Japan

Valvatidae
Small wide-spired operculate snails, commonly refered to as valve snails, egg-laying and
hermaphroditic. Burch (1982) lists 11 North American species. 11 are also listed for the former
USSR by Zhadin (1952) while only two are listed for India, and one for Africa. Banarescu
includes the following genera;
Valvata; (several subgenera listed by both Banarescu and Burch) throughout Europe, northern Asia, and North
America.
Borysthenia; Eastern and central Europe.
Gagea; Endemic in Lake Ohrid.

They have a featherlike gill, visible on the left side outside the shell when the snail is active
(Brown, 1991), and a ciliated pallial tentacle extending out to the right. Please see the Animal
Diversity Web http://animaldiversity.ummz.umich.edu/site/accounts/information/Valvatidae.html
or http://www.allesumdieschneck.de/html/valvatidae.html for pictures of living Valvata snails.
Dillon (2000) says of this little studied group, that some researchers have reported filter-feeding
behaviour, while others have concluded they are strictly grazers. Shell characteristics may be
variable. For instance the species V. tricarinata occurs in forms having differing numbers and
locations of spiral carinae or angulations. Several such forms may occur within a single
population (William Heard, in Burch, 1982).

Valvata stenotrema (Polinski) Lake Ohrid, Macedonia. x8.

Valvata bicarinata Lea 1841 American midwest and northeast. x8.

Megalovalvata demersa (Lindholm, 1909) Lake Baikal. x8.

Potamididae
A few members of this marine or brackish water family have adapted to fresh waters. Most inhabit marginally
marine environments in the tropics.

Telescopium telescopium
(Linnaeus, 1758) Brandt
(1974) indicates this is a snail
of "muddy irrigation trenches,
drainages and swamps in the
mud flats." Native to much of
Southeast Asia.

Cerithidea cingulata
(Gmelin, 1790) Native to
Thailand. This image,
and the photographs
below are from Brandt (1974).

Buccinidae
Carniverous gastropods related to the marine whelks. Two freshwater genera, Clea and
Afrocanidia.
Brandt (1974) describes and pictures six Clea (subgenus Anentome) species, but could only
confirm Clea helena for the territory of Thailand. He says of Clea helena that it is the only
species in Thailand that is "not restricted to running water as it is also found in lakes and ponds",
and that it "feeds predominantly on decaying protein, but has been observed to attack living
snails and worms."

Clea has been offered for sale in the pet trade and the possibility exists for it to become invasive.
Harry Lee includes this and other information on the genus at http://www.listserv.uga.edu/cgi-bin/wa?
A2=ind0712B&L=conch-l&P=R722&D=1&H=0&O=T&T=1. He also posted a listing of species at
http://www.listserv.uga.edu/cgi-bin/wa?A2=ind0712B&L=conch-l&P=R1895&D =1&H=0&O=T&T=1.

Clea helena (Philippi, 1847) (Left)

Widespread, Southeast Asia, Indonesia.
Variable. x3.

Clea scalarina (Deshayes, 1876)

(Right)
Mekong River. Slightly magnified.

Nassariidae
Nassa mudsnails, characteristic of tidal
mudflats, have also invaded fresh waters.
Banarescu (1990) mentions
Pygmaenassa - India,
Nassodonta - East Asia, and
Arcularia - Lake Chilka, Burma.

Nassodonta dorri
(Wattebled, 1886) Viet Nam.
Photo Bill Frank, webmaster
Jaxshells.org. x3.

Marginellidae

Rivomarginella morrisoni Brandt, 1968. x4

Rivers, lakes, and canals along the Gulf of Thailand. One
freshwater species "Known from Thailand only" (Brandt, 1974).

Pyramidellidae

Morrisonietta siamensis Brandt, 1968. x10

Prefers brackish water of low salinity. Several species endemic
to various localities in Southeast Asia (Brandt, 1974).

Freshwater Molluscan Shells / "Pleuroceridae"

Io fluvialis (Say, 1825) in habitat, northeast Tennessee, with several Leptoxis, in strong current.
There have been several attempts to re-introduce this species into suitable parts of its former range, with some
success.

"Pleuroceridae": (Cerithioid snails not in families Thiaridae or Melanopsidae)

Diverse in rivers of the Southeast United States. Many genera world wide. Burch in 1982 (note
p. 196) discusses the various names that are applied to this family and concludes "...there is as
yet no really solid basis for adequately comparing Pleurocera and its allies with Paludomus and
its related taxa or Pachychilus and its relatives. Until the necessary comparative studies have
been completed and evaluated, perhaps it is best to retain the family name Pleuroceridae."
Alternative classifications for the Pleuroceridae were considered by Banarescu (1990), who
finally settled with the subfamilies and genera in the table below. Since that time, many genera
have been reclassified, and several subfamilies have been raised to family status. Please see
Bouchet and Rocroi (2009).

Pleurocerinae
Pleurocera; American interior
Elimia; widespread, eastern North America
Lithasia; Ohio River drainage
Leptoxis; widespread, eastern North America
Io; endemic to Tennessee River and tributaries
Gyratoma; endemic to Coosa River
drainage, largely extinct
Juga; American far West and Northwest.
Holandriana; Danube River basin
Namruta; Japan
Amnipila; central Mexico
Pachychilus; Central and much of South America
Doryssa; eastern South America
Potadoma; central Africa
Potadomoides; southeastern Africa
Pachymelania; central Africa

Lavigeriinae
Semisulcospira; East Asia and Japan
Lavigeria; Lake Tanganyika

Paludominae
Paludomus; India and Southeast Asia
Chlorostracia; part of Burma
Cleopatra; Widespread Central Africa and Nile
Pseudocleopatra; tropical West Africa
additional genera endemic to Lake Tanganyika

Paramelaniinae = Tiphobiinae

Elimia catenaria (Say, 1822)

Atlantic coastal plain rivers

Tiphobia; Lake Tanganyika.

According to Dillon (2000), Pleuroceridae feed on detritus or the algae growing on hard surfaces,
and many seem to be able to feed on both. The family is characterized by having two sexes, male
and female, but with no outward expression of gender. Reproduction is iteroparous, meaning that
they may do so several times throughout their multi-year lifespans. Fertilization may occur by
males releasing sperm directly into the environment, as mating behaviour has never been
observed. Eggs are laid on a hard surface, for example the underside of stones, and develop into
tiny snails without a larval stage.

Lithasia snails with egg masses in a tributary of the Tennessee River, laid in early spring. slightly enlarged.
Photo by Mark Whited.

Syrnolopsidae, included for convenience with the African pleurocerids, but considered a
seperate family.
Three genera endemic to Lake Tanganyika.

To North American Pleuroceridae

To African Pleuroceridae (Paludomidae, Potamididae, Syrnolopsidae)

Latin American Pleuroceridae (Pachychilidae)

Pachychilus indiorum (Morelet, 1849),

Central America. This, along with P. glaphyrus,
are the edible "jute snails" of the Mayas. Hundreds
have been found in archaeological sites along with
the clam Nephronaias ortmanni. (website)
Kevin Cummings indicates these freshwater snails
are fairly widespread in Central America (Mexico,
Honduras, Belize, Guatemala), and Northern South
America (Venezuela). There are at least 111 described
taxa, many of which are recognized as synonyms.

Doryssa pernambucensis
(Reeve, 1861), Brazil

Doryssa cachoeirae F. Baker, 1913

Brazil

Asian Cerithioid snails (Paludomidae and Semisulcospiridae)

Paludomus chilinoides Reeve 1847. Sri Lanka rivers. Preston (1914) lists Paludomus neritoides
64 species and a number of varieties in four subgenera: typical Paludomus, (34
species); Philopotamus (9 species) Tanalia (20 species) and Stomatodon (1
species). Most are native to Sri Lanka, with some living elsewhere in Southern
and Southeast Asia. Brandt (1974) only lists one additional species for the
territory of Thailand. x4

Reeve 1847. Sri Lanka.

Approx. natural size.
Large, thick shell, patterns
visible when backlit. In
subgenus Tanalia. x1.

Paludomus loricata (Reeve). This species is sold in Europe as an aquarium snail. Listed by
Preston, (1914) in the subgenus Tanalia, native to Sri Lanka "delighting in the most rapid
mountain torrents." Photo by Eduardo Diaz, Spain.

Paludomus regalis Layard, 1854. Sri Lanka. x 1.5.

Balanocochlis glandiformis (Schepman, 1896), and operculum, Malaysia.

Semisulcospira libertina
(Gould, 1858) Taiwan

Semisulcospira
reticulata
Kajiyama and Habe,
Japan

Biwamelania japonica (Smith, 1876)

Lake Biwa, Japan

Freshwater Molluscan Shells

Thiaridae
The general range of the thiaridae is circum-tropical, and many of the species are extremely widespread. Their dispersal
is facilitated by the fact that all are parthenogenic females. Melanoides tuberculata (Mller 1774), similar to M.
maculata, below, is a common aquarium snail. Among the genera are the following;
Thiara; East Africa to Polynesia, some species also very wide ranging.
Melanoides; Southern Europe, Africa
Brotia; Southeast Asia
Terebia; south Asia, Oceania
Sermyla; southeast Asia, Oceania
Sermylasma; Australia
Pseudopotamis; Australia
Melanatria; Madagascar
Fijidoma; Fiji Islands
Tylomelania; Celebes
Cubaedomus; Cuba
Aylacostoma; Central and South America
Doryssa; South America (see Pleuroceridae)
Hemisinus; South America

Australia

Melanoides denisoniensis Thiara balonnensis (Conrad, Syrmylasma venustula (Brot, 1877),

Queensland, Australia
(Brot, 1877), Queensland,
1850), Queensland, Australia.
Australia

Southeast Asia

Paracrostoma
pseudosulcospira
armata (Brandt, 1968)

Brotia pagodula (Gould, 1847)

Brotia henriettae
(Gray, 1834)

An internet search for Brotia pagodula and B. henriettae yields a number of German aquariist sites. It appears that there
is a substantial trade in several Southeast Asian species to supply that market as of 2008. Shell characteristics suggest

the specimens pictured here were part of that trade, and that they may have matured in their natural environment prior to
being collected and shipped off. http://www.allesumdieschneck.de/html/brotia_pagodula_english.html (accessed Sept.
2008) indicates that Brotia pagodula requires constantly moving highly oxygenated water and that most die after 6
months in an aquarium. If they are not bred in captivity it is probably best to avoid them.

Geoff Macaulay provides the following information regarding Brotia costula.

Brotia (Brotia) costula (Rafinesque, l833), is a large and widespread snail in the family Thiaridae (?) [Banarescu does not mention Brotia in his
discussion of either Thiaridae or Pleuroceridae.] The best description I have is in Archiv fur Molluskenkunde 105, I-IV, 1-423, The Non Marine
Aquatic Mollusca of Thailand by Rolf A. Brandt (1974). To briefly summarise this - This is an extremely variable species and also the largest
Brotia. There are a great many synonyms and a wide distribution from North India, Burma, South China, continental South East Asia to
Sumatra, Java and Borneo. It occupies many different habitats. Brandt recognises 3 races: Brotia costula costula (Rafinesque 1833), B. c
varicosa (Troschel 1837) and B. c peninsularis Brandt l974. Synonyms for costula costula include variabilis Benson, herculea Gould, indica
Souleyet, corrugata Reeve, jullieni Deshayes, and peguensis Morelet.

Also see Khler and Glaubrecht's 2006 paper on the systematics of Brotia, placing the genus in the family
Pachychilidae: http://download.naturkundemuseum-berlin.de/frank.koehler/Brotia_2006.pdf

Indo-Pacific

Thiara winteri Von Dem Busch, 1842, Thiara, unknown sp., Thiara, unknown sp., Philippines. Small
Philippines, x2

Philippines, x2.

shell, x4.

Adult, front and back, and immature individuals, more magnified. Unknown Thiara species,
probably T. amarula, T. terpischore, or T. macrospira. Photos and ID's provided by Don Barclay.

Melanoides torulosa
(Bruguiere, 1789)
Philippines

Unknown species, Philippines

Unknown Melanoides sp.

Philippines

Melanoides maculata
(Bruguiere, 1789), Philippines.

Thiara cancellata, (Roeding, 1798)

cleaned with bleach, from
the Philippines. Projections are shelly.

Melanoides granifera, (Lamarck, 1822) Philippines.

South America

Another presumptive Thiara cancellata(?) Java

rivers. The thin projections are periostracal (soft organic)
material.

Hemisinus eduardsi Aylacostoma pulcher Aylacostoma francana

(Lea, 1852), Brazil, (Reeve, 1860), Brazil, (Ihering, 1909). Sao
slightly enlarged.
slightly enlarged.
Paulo State, Brazil. x3.

Africa
Pilsbry and Bequaert (1927) list 23 species and 8 additional subspecies of Melanoides for the African continent outside
Lake Nyassa. They include an additional list of 38 species (in several dubious genera) for that lake compiled from
Bourguignat, but add "We do not care to add new name combinations for a host of merely mutational forms" They felt
the task of determining the actual number of species in Lake Nyassa, taking into account variation typical within
species, was outside the scope of their work. Below is a small sample of those figured from the rest of the continent.
Left to right:
Melanoides anomala (Dautzenberg and Germain, 1914) 8 mm
M.liebrechtsi (Dautzenberg, 1901) 24 mm.
M. bavayi (Dautzenberg and Germain, 1919) ? mm
M. nsendweensis consobrina (Dupuis and Putzeys 1900) 13.5 m

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Freshwater Molluscan Shells

Thiaridae
The general range of the thiaridae is circum-tropical, and many of the species are extremely
widespread. Their dispersal is facilitated by the fact that all are parthenogenic females.
Melanoides tuberculata (Mller 1774), similar to M. maculata, below, is a common aquarium
snail. Among the genera are the following;
Thiara; East Africa to Polynesia, some species also very wide ranging.
Melanoides; Southern Europe, Africa
Brotia; Southeast Asia
Terebia; south Asia, Oceania
Sermyla; southeast Asia, Oceania
Sermylasma; Australia
Pseudopotamis; Australia
Melanatria; Madagascar
Fijidoma; Fiji Islands
Tylomelania; Celebes
Cubaedomus; Cuba
Aylacostoma; Central and South America
Doryssa; South America (see Pleuroceridae)
Hemisinus; South America

Australia

Melanoides

Thiara balonnensis (Conrad, Syrmylasma venustula (Brot, 1877),

denisoniensis
1850), Queensland, Australia.
(Brot, 1877), Queensland,

Queensland, Australia

Australia

Southeast Asia

Paracrostoma
pseudosulcospira
armata (Brandt,
1968)

Brotia pagodula (Gould, 1847)

Brotia
henriettae
(Gray, 1834)

An internet search for Brotia pagodula and B. henriettae yields a number of German aquariist
sites. It appears that there is a substantial trade in several Southeast Asian species to supply that
market as of 2008. Shell characteristics suggest the specimens pictured here were part of that
trade, and that they may have matured in their natural environment prior to being collected and
shipped off. http://www.allesumdieschneck.de/html/brotia_pagodula_english.html (accessed
Sept. 2008) indicates that Brotia pagodula requires constantly moving highly oxygenated water
and that most die after 6 months in an aquarium. If they are not bred in captivity it is probably
best to avoid them.

Two specimens obtained from diferent sources, identified as

Brotia costula (Rafinesque, 1833), (left), and Brotia
costulata (Rafinesque, 1833), (right), from Malaysia, and
Bangladesh, respectively. Both of these morphologies fall within
the very variable subspecies Brotia (Brotia) costula costula.

Brotia insolita (Brot, 1868)

Cambodian rivers. Brandt (1974)
lists ten species and four additional
subspecies within the genus
Brotia.

Geoff Macaulay provides the following information regarding Brotia costula.

Brotia (Brotia) costula (Rafinesque, l833), is a large and widespread snail in the family Thiaridae (?) [Banarescu
does not mention Brotia in his discussion of either Thiaridae or Pleuroceridae.] The best description I have is in
Archiv fur Molluskenkunde 105, I-IV, 1-423, The Non Marine Aquatic Mollusca of Thailand by Rolf A. Brandt
(1974). To briefly summarise this - This is an extremely variable species and also the largest Brotia. There are a
great many synonyms and a wide distribution from North India, Burma, South China, continental South East Asia to
Sumatra, Java and Borneo. It occupies many different habitats. Brandt recognises 3 races: Brotia costula costula
(Rafinesque 1833), B. c varicosa (Troschel 1837) and B. c peninsularis Brandt l974. Synonyms for costula costula
include variabilis Benson, herculea Gould, indica Souleyet, corrugata Reeve, jullieni Deshayes, and peguensis
Morelet.

Also see Khler and Glaubrecht's 2006 paper on the systematics of Brotia, placing the genus in
the family Pachychilidae: http://download.naturkundemuseumberlin.de/frank.koehler/Brotia_2006.pdf

Indo-Pacific

Thiara winteri Von Dem Busch, 1842, Thiara, unknown

Philippines, x2
sp.,
Philippines, x2.

Thiara, unknown sp., Philippines. Small

shell, x4.

Adult, front and back, and immature individuals, more magnified. Unknown Thiara species,
probably T. amarula, T. terpischore, or T. macrospira. Photos and ID's provided by Don Barclay.

Melanoides torulosa
(Bruguiere, 1789)
Philippines

Unknown species, Philippines

Unknown Melanoides sp.

Philippines

Melanoides maculata
(Bruguiere, 1789), Philippines.

Thiara cancellata, (Roeding, 1798)

cleaned with bleach, from
the Philippines. Projections are shelly.

Another presumptive Thiara cancellata(?)

Melanoides granifera, (Lamarck, 1822)
Philippines.

South America

Java
rivers. The thin projections are periostracal (soft
organic)
material.

Hemisinus eduardsi Aylacostoma pulcher Aylacostoma francana

(Lea, 1852), Brazil, (Reeve, 1860), Brazil, (Ihering, 1909). Sao
slightly enlarged.
slightly enlarged.
Paulo State, Brazil. x3.

Africa
Pilsbry and Bequaert (1927) list 23 species and 8 additional subspecies of Melanoides for the
African continent outside Lake Nyassa. They include an additional list of 38 species (in several
dubious genera) for that lake compiled from Bourguignat, but add "We do not care to add new
name combinations for a host of merely mutational forms" They felt the task of determining the
actual number of species in Lake Nyassa, taking into account variation typical within species,
was outside the scope of their work. Below is a small sample of those figured from the rest of the
continent.
Left to right:
Melanoides anomala (Dautzenberg and Germain, 1914) 8 mm
M.liebrechtsi (Dautzenberg, 1901) 24 mm.
M. bavayi (Dautzenberg and Germain, 1919) ? mm
M. nsendweensis consobrina (Dupuis and Putzeys 1900) 13.5
m

Freshwater Molluscan Shells

Melanopsidae

The Melanopsidae have a strange disjunct distribution, mostly around the Mediterranean.
Melanopsis is native to Spain, Morocco, and the Middle East,
Esperiana is found in Central Europe, but their closest relative,
Zemelanopsis, is native to New Zealand and New Caledonia.
Faunus (placed in the family Thiaridae by many authors, and Potamididae by Brandt (1974))
ranges through Southeast Asia, the East Indies, and Madagasgar.
According to Behrendt, the Melanopsidae can feed on detritus or the algae growing on hard
surfaces, and many are willing to consume either. Sexes are separarate, and adult females have a
small whitish reproductive opening on the right side of the foot. Reproduction occurs several
times throughout the multi-year life of the snail. Fertilization may occur by males releasing
sperm directly into the environment, as mating behaviour has not been observed. Eggs are
usually layed in clutches on hard surfaces such as the underside of stones, but may also be
scattered on soft substrates, and hatch into tiny snails without going through a larval stage.

Northern Africa

Melanopsis attenuata (Pallary,

1920),
Morocco, from two different sources.
The
magnified specimen (left) shows no
trace of the
color banding seen in the specimens
above.

Melanopsis cesari Pallary,

1920.
Morocco. x3.

Melanopsis buccinoidea Olivier, Jordan

Melanopsis foleyi (Pallary),

River

Morocco

Melanopsis magnifica Bourguignat,

with periostracum removed, Morocco.

Melanopsis magnifica ingens Pallary, Morocco.

These specimens were obtained from dealers identified as "Melanopsis praemorsa (Linnaeus,
1758)", the left pair from Crete, Greece, and the right from the Sea of Galilee. Van Damme (1984) also includes
within this named species all of the forms from northern Africa, some of which are shown above. This may have
been in response to the earlier excesses of Jules Rene Bourguignat, "one of the epigons of Victorian species-makers,"
Van Damme expressed the opinion that his species splitting, ignoring natural variation, caused permanent damage to
African malacology. He goes on to mention that Pallary was "a staunch devotee of Bourguignat."

Europe

Esperiana esperi (A. Ferrusac, 1823), Danube Esperiana daudebartii Unknown Melanopsis sp
River,
Hungary.

acicularis (Prevost
1821),

Hot spring in Zaragosa, Spain.

Romania

Melanopsis bicarinata Spain

Indo-Pacific Faunus species

Faunus ater (Born, 1778), native to Philippines, Southeast Asia, and the

unknown Faunus species,

East
Indies. Brandt (1974) includes Faunus with the family Potamididae and
indicates
that "It lives in fresh as well as slightly brackish water near the coast in creeks,
small rivers and lagoons."

Madagascar.

New Zealand and Oceania

Zemelanopsis trifasciata Gray, 1843,

North Island of New Zealand, streams.

Melanopsis frustulum
Morelet, 1856. New Caledonia

Freshwater Molluscan Shells

Hydrobiidae (Somatogyrus), unknown species, from the Chunkey River, Mississippi, United
States. Collected September 2000.

Hydrobiidae and Allied Families (Superfamily Rissoacea,

Hydrobioidea, or Truncatelloidea)
World wide, these are the small ones, ranging down almost to the microscopic. Magnifications
used here are approximately 4 x and 8 x, for filenames ending with 300 and 600 respectively (see
scales). In the author's experience in the south-central United States, such "microshells" are
generally uncommon, only occasionally found in flood-deposited creek drift. In some instances
they can be abundant, as in widely scattered springs, or at one locality in coastal Georgia where a
few seeps flow around cement sidewalks, and another, a reach of the Chunkey River in
Mississippi, where they swarmed over bedrock, filamentous algae, and sand. Thompson, in
http://www. flmnh. ufl.edu/natsci/malacology/fl- snail/snails1.htm indicates that many species
are annual, with a single brood, and a complete turnover of individuals every year. He also points
out that there is a great deal more variation between various taxa than would be suggested by the
often similar shells.

Classification:
The classification of this enormous group of organisms is under review. Banarescu provides the
following information:
Pomatiopsidae: Two subfamilies Pomatiopsinae, and Triculinae. Pomatiopsinae contains genera Erhaia,
China; Aquidauania, central South America; Tomichia, South African rivers; Coxiella and Coxielladda from west
and south Australia; Oncomelania, from east Asia; Blanfordia, and Fukuia, Japan; Cecina, Japan, Manchuria, and
western North America; and Pomatiopsis, North America. The Triculinae include twelve genera, all in south and
southeast Asia. Nine are endemic to the Mekong River.

Bithyniidae: Gabbiella, central Africa; Jubaia, and Incertihydrobia, east Africa; Sierraia, west Africa;
Funduella, Limnitesta, and Congodoma, south-central Africa; Gabbia, central Australia; Emmericiopsis, Mysorella,
Parabithynia, Pseudovivipara, and Sataria, endemic in various locations, south, and southeast Asia; Hydrobioides,
south Asia; Petroglyphus, Phillipines; Allocinma, widespread in southern Asia; Bithynia, Europe through central
Asia, introduced (possibly native) to North America.

Baicaliidae: Baicalia, endemic to Lake Baical.

Benedictiidae: Kobeltocochlea, Lake Baical and Lake Hubsugul (Mongolia); Benedictia, Lake Baical.
Pyrgulidae: Several genera in southern Europe and Turkey, one, Antroselates, in Indiana and Kentucky caves.
Burch, 1982 includes this later with family Micromelaniidae.

Hydrobiidae: A very large family with hundreds of genera. These can be grouped into a number of subdivisions,
many of which are considered distinct families or subfamilies by various authors. Banarescu discusses some of the
disagreements in classification schemes, and offers the following informal groupings, or "lineages":
Lithoglyphine, eastern and far western North America, southern Europe.
Glacidorbis, Tasmania, south Australia, and Chile.*
Orientaliine - horatiine - amnicoline, eastern and north-central North America; many genera, subfamilies in southern
and central Europe, Turkey.
Fontigentine, eastern North America.
Nymphophiline; most of area of United States.
Cochliopinine; Central America, Antilles, Peru.
Littoridinine; Mexico, Central and northern South America.
Mexithaumine; Rio Grande, Equador.
Emmericiine; western Balkan Peninsula.
Lithoglyphuline; western Balkan Peninsula, Danube basin.
Bythinelline; Europe, largely on southern peninsulas.
Moitessierine; Europe, largely on southern peninsulas.
Fluvipupa; Australia, New Guinea, Tasmania, New Zealand.
* Ponder (1986) assigns the genus Glacidorbis to its own family and superfamily within the pulmonate
Basommatophora!

A commonly used classification scheme for the hydrobiidae is based on anatomical structures of
the male reproductive organ or "verge" (Burch,1982), although there is question as to whether
the resulting system is correct. He includes three families, five subfamilies, 29 genera, four
additional subgenera, and 158 species for North America in his superfamily Truncatelloidea. He
places the family Bithyniidae, with its one probable introduced species in North America, in the
superfamily Ampullarioidea, (Viviparoidea) however, with the much larger Viviparidae and
Ampullariidae.
Truncatelloidea
Micromellaniidae: Antroselates spiralis alone in North America is included in this family.
Based on radular characteristics, it includes species from Lake Baikal, and other large lakes in
southeastern Europe and Asia.
Hydrobiidae:
Hydrobiinae: Aphastracon, Hoyia, hyalopyrgus, Littoridinops, Probythinella, Pyrgophorus, Tryonia.
Lithoglyphinae: Antrobia, clappia, Cochliopina, Fluminicola, Gillia, Lepyrium, Somatogyrus (Somatogyrus

s.s., and Walkerilla).

Nymphophalinae: Birgella, Cincinnattia, Fonticella (Fonticella s.s., Microamnicola, and Natricola),

Marstonia, Notogilla, Orygoceras (name under review), Pyrgulopsis, Rhapinema, Spilochlamys, Stiobia..
Amnicolinae: Amnicola (Amnicola s.s., and Lyogyrus), Hauffenia, Horatia.
Fontigentinae: Fontigens.
incertae sedis: three species.
Pomatiiopsidae Pomatiopsis.

North America

Pomatiopsis lapidaria (Say, 1817)

Fluminicola nuttalliana (Lea, 1838)

south central United States. (Pomatiopsidae)

Columbia River basin, Pacific northwest.

(Hydrobiidae) Up to 10 mm, relatively large
for the family, it inhabits most creeks and
small rivers. Operculum, inner side, upper
left.

Probable Marstonia species, from spring in

upper East Tennessee, February, 2002. Of a
number of specimens examined, none displayed

the verge, suggesting parthenogenicity in the

population.

Europe

Chilopergula prespensis
(Urbansky) Macedonia.

Emmercia patula (Brumati, 1838)

Bithynia leachi Italy.

Croatia. (Micromelaniidae)

(Bithyniidae)

(Micromelaniidae)

Lithoglyphus naticoides (Pfeiffer)

Potamopyrgus jenkensi E. A. Smith

Hungary. (Hydrobiidae)

Hungary. (Hydrobiidae)

Asia

Bythnia kiusiuensis (Hirase,

1957)

Gabbia longicornis (Benson)

Viet Nam. (Bithyniidae)

Japan. (Bithyniidae)

Bithynia tentaculata (L.,

1758)
Europe, Asia, and North America
(introduced). (Bithyniidae)

Stenothyra basiangulata Mori,

1838 Japan. (Stenothyridae) Brandt

Caspiohydrobia issykkulensis
(Clessin, 1894) Kyrgystan.

(1974) includes 16 freshwater and 15

brackish water species for the territory
of Thailand in Stenothyra.

(Hydrobiidae)

Hydrobioides striatulus
(Benson) Viet Nam. (Hydrobiidae)

Blanfordia japonica
(A. Adams, 1854)

Fluvicingula elegantula
(A. Adams, 1861), Japan.

Japan. (Pomatiopsidae)

(Iravadiidae)

Paludinella devilis
(Gould, 1861) Japan.
(Hydrobiidae)

Mekong River Pomatiopsidae:

Paraprosthenia
levayi (Bavay,
1895) x8

Lacunopsis harmandi
Poirier, 1881. x4

Pachyhydrobia
spinosa Poirier,
1881 x4 Note
single spine on
back of shell.

to Australian region

South America

Litoridina australis

Potamolithus jacuyensis Pilsbry,

Jullienia harmandi
Poirier, 1881.
x4

(d'Orbigny, 1835),

1899 Brazil. (Hydrobiidae)

Uruguay. (Hydrobiidae)

Freshwater Molluscan Shells / Pulmonata / Basommatophora

Physella sp.

Freshwater Pulmonate Gastropods

Freshwater pulmonate snails are characteristically thin and light shelled. Many can regulate their
buoyancy to neutral and even positive by taking air into a "lung," which allows them to glide
upside down on the air/water surface. Their life cycle is rapid. In nature, one or two generations
per year may be produced, with complete replacement of individuals (Brown, 1991). All are
hermaphroditic; that is, each individual has the capacity to function as both male and female.
Unlike the prosobranchs, with many endemic species and varieties, the pulmonate species are
fewer but much more widespread. Freshwater pulmonates reach their greatest size and diversity
in the northern latitudes, and many species are commonly found in shifting and ephemeral
habitats.
Although all are in the subclass Pulmonata, the freshwater pulmonate snails are not closely
related to the land snails. Terrestrial pulmonates ("true" land snails) are in the Order
Stylommatophora, while the freshwater pulmonates, considered as the single superfamily
Lymnaeacea by some authors, are placed in the Order Basommatophora, along with the marine
pulmonate "false limpets," superfamily Siphonariaceae. Freshwater limpets are also
Lymnaeacean, but their limpet-like form has evolved several times within this group, from
different coiled ancestors. The marine superfamily Melampoidea (Ellobioidea) are included as a
third grouping within Basommatophora by some authors, but are given a separate order,
Archaeopulmonata, in more recent sources.
Abbott (1989) uses:

Order Archeopulmonata, with a single superfamily,

Ellobioidea (littoral marine "melampus" shells)
Order Basommatophora, with seven superfamilies:
Amphiboloidea (marine)
Chilinoidea
Chilinidae (South American freshwater snails)
Latiidae (New Zealand freshwater limpets)
Siphonarioidea (marine "false limpets")
Acroloxoidea (freshwater limpets [in part])
Lymnoidea (dextral* pond snails, limpets)
Physoidea (sinistral* pond snails)
Planorboidea (ramshorn pond snails, limpets).
Banarescu (1990) for freshwater taxa only, uses:
Order Basommatophora with a single freshwater superfamily;
Lymnaeacea, containing six families in two sublineages:
Chilinidae (South American freshwater snails)
Latiidae (New Zealand freshwater limpets)
Acroloxidae (freshwater limpets [in part]), and
Lymnaeidae (dextral* pond snails, some freshwater limpets)
Physidae (sinistral pond snails)*
Planorbidae (ramshorn pond snails, other freshwater limpets)
* Sinistral and dextral refer to the mapping of rotation direction to a straight line vector. Most
snail shells, as well as common screws, etc., are dextral, or right-handed. Take your right hand.
Point the thumb in the general direction the screw moves as it rotates or the shell grows as it
spirals downwards (down), and your curled fingers will define the direction of rotation for righthanded shells, screws, etc. Likewise for sinistral or left-handed snail shells, but in reverse.

Freshwater Molluscan Shells: Pulmonata

families [Physidae] [Lymnaeidae] [Chilinidae] [Planorbidae]

Lymnaeidae
The Lymnaeidae are cosmopolitan, with members on all continents and large islands, and even
some smaller oceanic islands. Most, in the subfamily Lymnaeinae, are dextral pond snails, while
those in the subfamily Lancinae are limpet-like. Burch lists 57 species in nine genera for North
America. In common with other freshwater pulmonates, all feed on algae and detritus, and all are
hermaphroditic. Species are fewer and smaller in the tropics, where they may carry parasites
including trematodes and flukes.
Burch, (1982) uses the following classification for North American Lymnaeids:

Family Lymnaeidae
Subfamily Lymnaeinae
Genus Acella
Genus Bulimnea
Genus Fossaria, with subgenera Fossaria ss., and Bakerilymnaea
Genus Lymnaea
Genus Pseudosuccinea
Genus Radix
Genus Stagnicola, with subgenera Stagnicola ss., and Hinkleyia

Subfamily Lancinae (all limpet-like; see Freshwater limpets)

Genus Fisherola
Genus Lanx, with subgenera Lanx ss, and Walkerola

Bulimnea megasoma (Say, 1824) North-central United States and adjacent Canada, shallow water, smaller
lakes.

These two species are commonly found

crawling on mud and debris, just above the
waterline of ponds or slow-moving creeks.

Fossaria obrussa Say, 1825. Southeastern United

States.

Pseudosuccinea columela (Say, 1817)

Eastern North America

Stagnicola elodes
(Say, 1821) Much of
the
U. S. and Canada.

Stagnicola caperata (Say,

1829) Northern North America.

Stagnicola
montanensis
(Baker, 1913)
Rocky Mountains.

Stagnicola emarginata
(Say, 1832)
Eastern Canada and Northeast
U.S.

Lymnaea peregra (Muller, 1774)

Europe. Halstead's Golden Guide to
Tropical Fish indicates this species
should be avoided in aquaria because
it produces a poison fatal to fish.

Lymnaea stagnalis (Linne, 1758) Europe, also North America.

Lymnaea luteola australis Annandale &

Rao,
1925. Sri Lanka. x3.

Lymnaea (Stagnicola) fuscus (C. Pfeiffer, 1821) from

Bulgaria
(left) and Lymnaea
Kazakhstan (right).

palustris (Muller, 1774), from

Identifications provided by Richard Ottermans. Both x2.

Pseudosuccinea
peregrina (Clessin,
1822) Farm ponds,
southern Brazil

Austropeplea lessoni (Deshayes, 1830) New South Wales,

Australia.

Lymnaea (Radix) natalensis Krauss, 1848

South Africa. Image from Pilsbry and Bequaert, 1927,
courtesy American Museum of Natural History

Freshwater Molluscan Shells: Pulmonata / Physidae

families [Lymnaeidae], [Planorbidae], [Chilinidae]

The Physidae, sinistral pond snails, are commonly referred to as tadpole snails or pouch snails.
They are widespread, abundant, and tolerant of pollution. Burch (1982) indicates that they are
most abundant in the New World. They have evidently found a shell morphology suitable for
their life station, as he goes on to say "...the physids have undergone considerable diversification,
much of which is not clearly exhibited in their shells. Many of the species [and genera] are not
easy to identify on shell characters alone."

They have been used in studies of ecophenotypic plasticity, a so-called phenoplastic switch. Burt
Vaughan of Washington State University indicates several studies in M. J. West-Eberhardt's
recent compendium of research, "Developmental Plasticity & Evolution" (Oxford Press, 2003,
pp. 307-362). A typical example involved rearing Physa gyrina, or P. heterostropha in controlled
pair groups in either water in which crayfish co-existed or water in which only fish co-existed.
Within a month, differences in shell morphology appeared; i.e., snails exposed to shell-crushing
fish predators showed wide apertures and very much strengthened, rotund shells. Snails exposed
to crayfish only showed narrow-apertured, thin elongate shells, with barricading teeth.
Banarescu lists the following classification for the Physidae:

Family Physidae (with number of taxa listed by Burch (1982) for North America:)
Subfamily Physinae
Genus Physa most of Northern Hemisphere. (2 sp. plus 1 additional morph)
Genus Physella, Holarctic,containing subgenera;
Physella s.s. (16 species, 15 subspecies, and several additional morphs), subgenus
Costatella, (14 species with 13 additional subspecies and morphs, and subgenus
Petrophysa (1 species).

Subfamily Aplexinae
Genus Aplexa most of Northern Hemisphere. (1 species plus 1 morph)
Genus Stenophysa Central America Mexico, introduced in Texas. (2 species)

Physella species, probably Physella

gyrina

Physella heterostropha halei Unidentified Physella

(Lea, 1864) Central and Southern species,

aurea (Lea, 1838). East and central Unite

United States, ponds.

small western Montana creek.

States. Many subspecies and morphs of

Physella
gyrina have been named.

Aplexa hypnorum, Moscow, Russia

Physa albertisi Clessin,

1866. Papua New Guinea

Physa cubensis
Brazil

Freshwater Molluscan Shells: Pulmonata

families [Physidae], [Lymnaeidae], [Chilinidae]

Family Planorbidae
Generally referred to as "Wheel Snails", "Orb Snails", or "Ramshorn snails", the Planorbidae
are the largest family of aquatic pulmonate gastropods, with species present on all continents and
most islands. They are sinistral in their orientation, in spite of the fact that some may appear
dextral due to the spire being sunken more than the umbilicus. Many of these species harbor the
larvae of parasitic worms, particularly southern and Old-World taxa. Some are patelliform or
limpet-like (see Freshwater limpets). Banarescu (1990) provides the following classification,

here tabulated. Many subgeneric and subspecific names are in use, so for instance, the first
specimens pictured are Planorbella (Pierosoma) trivolvis trivolvis (Say, 1817).

Subfamily

Tribe

Rhodacmeinae

Rhodacmea; Southeast North America. Limpet-like.

Bulinini

Bulininae

Genus

Physastrini

Camproceratini

Bulinus; Africa, West Asia, southern Europe

Indoplanorbis; Southern Asia
Laevapex; eastern North America, limpet-like.
Gundlachia; South and Central America
Isidorella; south and west Australia
Bayardella; localities in Australia
Patelloplanorbis; New Guinea, Limpet-like?
Oppletora; northern Australia
Ancylastrum; Tasmania Limpet-like?
Miratesta; Celebes Is.
Protancylus; Celebes Is. Limpet-like?
Pettancylus; Southeast Asia, Australia, and the Pacific, East
Africa. Limpet-like?
Barnupia; north Australia and New Guinea, East Africa,
Limpet-like
Ferrisia; temperate North America. Limpet-like
Glyptophysa; southern Australia, New Zealand
Physastra; Australia and New Zealand, New Guinea
Ameriana; North and northeast Australia
Helisoma North America
Planorbarius Siberia
Camptoceras east and Southeast Asia

Planorbulini

Planorbula, (including Menetus, Promenetus, and Planorbella

of Burch, (1982); all North American.

Biomphalarinii

Biomphalaria; central Africa and Central and South America

Drepanotrema; South America, Antilles

Planorbini

Planorbis; Europe, western Asia, northern Africa

Afrogyrus and Afrogyrorbis; East Africa and Madagascar
Anisus; Europe and West Asia.
Bathyomphalus; Europe and much of Siberia
Choanomphalus; Lake Baikal
Gyraulis; all of both Europe and Africa, except Sahara

Segmentini

Segmentina; most of Europe and Siberia

Hippeutis; most of Europe through central Siberia
Lentorbis and Segmentorbis; Central and Southern Africa
Polypylis; coastal eastern asia and Japan
Helicorbis; East and Southeast Asia

Ancylini

Ancylus; Northern Hemisphere, discontinuous, Limpet-like

Planorbinae

Uncertain

Neoplanorbis and Amphigyra; southeastern North America

Planorbella trivolvis (Say, 1817) Eastern and Central Canada and Northern U.S. Four views of two specimens:
1) Apertural view with left side to left. 2) Left or spire side of shell. 3&4) Right, or umbilical side

Planorbella scalaris
(Jay, 1839) Florida

Helisoma anceps
(Menke, 1830)

Planorbella campanula
(Say, 1821)

Eastern, central, and

northern North America.

North-Central North America

Planorbella tenuis (Dunker, 1850)

Biomphalaria havanensis (?) (Pfeiffer, 1839)

Southern United States, Mexico; ponds.

Southern U. S., Central America, West Indies; ponds.

Snail shell obtained from a breeder in Belgium, courtesy of

Gerald Depaus. Possibly Planorbella occidentalis

(Cooper, 1870) or Planorbella ammon (Gould, 1855)

Drepanotrema lucidum (Pfeiffer)

Anisus filiaris (Gredler, 1885)

West Indies

western Asia

Anisus vortex Europe

Bathyomphalis contortus

(Linne, 1758) Europe

Hippeutis umbilicalis
(Benson, 1836) x4 (left)
Gyraulis siamensis
(Martens, 1867) x4 (right)
Brandt (1974) lists six Gyraulis, one
Hippeutis and three Segmentina
species for the territory of Thailand,
some of which are hosts to
Echinostoma and other parasites.
All are small inconspicuous snails.
Images from Brandt (1974).

Anisus leucostoma
(Millet, 1813).
Hungary. x 6

Planorbis corneus (L., 1758) Europe

Planorbid species from a pond in

the
south-Central United States,
possibly an
aquarium escapee.

Indoplanorbis exustus
(Deshayes, 1834). Sri
Lanka. x3

Biomphilaria
tenagophila tenagophila
(Orbigny, 1835). Bahia
State, Brazil. x 3.

Isidorella hainesii (Tryon, 1866) Victoria, Australia.

Isidorella newcombi (Adams & Angas, 1864)

South Australia.

Bulinus senegalensis O. F. Mller,

1781

Glyptophysa gibbosa (Gould, 1546)

Queensland, Australia.

Africa. Largest specimen is 4.4 mm. Despite

the
resemblance of these shells to the Physidae,
they
have been included with the Planorbidae in all
sources available.
The name Bulinus was originally applied by
Mller to include Physa snails, and
controversy
has existed as to whether "Bulinus" might have
priority for Physa. (Pilsbry and Becquaert,
1927)

Physopsis
africana
globosa (Morelet,
1866). Africa, to
20 mm. long.
Black and white
images from
Pilsbry
and Becquaert,
1927, courtesy of
the American
Museum of
Natural
History

Freshwater Molluscan Shells

Chilinidae native only to the temperate southern regions of South America, one genus,
Chilina, with 23 species listed by Pilsbry (1911) for the area east of the Andes.
Pilsbry goes on to indicate
"Chilina occupies the temperate and cold zones of South America from the Tropic of Capricorn
to Cape Horn [and the Falkland Islands] ...Within their area, the Chilinidae are abundant snails in
all suitable stations... They swarm in springs, small streams, lakes, and in some places the
margins of rivers. They are most abundant southward, becoming rarer and local toward the
northern borders of their range. The species from west of the Andes are in all cases, so far as we
know, distinct from those east ot the divide....
The eastern fauna ... inhabits a comparitively arid region, poorly watered by roughly parallel
streams flowing southeastward into the Atlantic. Each of the principal river systems has its own
series of freshwater mollusks, in large part distinct specifically or racially from those of other
rivers."
Brace, (1983) indicates that they are adapted through both physical and behaviorial features to
burrowing in soft substrates . He also suggests that morphological evidence supports the

relationship of Chilina, and by inferance, the rest of the Basommatophorans, to primitive

opistobranch gastropods.

Chilina fluminea Orbigny, Buenos Aires Province, Argentina

Chilina dombeiana (Bruiguiere, 1789),

Rio Negro province, Argentina

Chilina globosa Sowerby Chilina neuquenensis

Marshall, 1933 Rio
1841. Buenos Aires Province, Negro
province, Argentina

Argentina. x3.

[Lymnaeidae], [Planorbidae], [Chilinidae] [Physidae]

[Home Page] [Site Ma

Freshwater Molluscan Shells: Limpets

families [Physidae] [Lymnaeidae] [Chilinidae] [Planorbidae]

Freshwater limpets are small to minute and, like freshwater mollusks in general, polyphyletic.
None are related to marine limpets. In freshwater mollusks, a limpet-like form has arisen
multiple times from more than one lineage of coiled aquatic pulmonate snails, but from
pulmonates alone. Somewhat similar forms from the family Latiidae (also pulmonate) and the
non-related Neritidae lack the centralized apex. The Lancinae are confined to the Pacific
drainages of North America while the Acroloxidae have an extremely disjunct distribution,
several widely separated localities scattered over the northern hemisphere. They share a dextral
organization of soft anatomy which is reflected in the assymetry of their shells. "Ancylidae"
(taxomomic groups within the Planorbidae) have a sinistral organization, and a corresponding
sinistral assymetry in the shell, and a near worldwide distribution.
Burch, (1982) uses the following classification for North American freshwater limpets:

Order

Superfamily

Family

Acroloxoidea

Acroloxidae

Acroloxus

Lymnaeidae Lancinae

Fisherola
Lanx, with subgenera
Lanx ss, and Walkerola

Limnophila Lymnaeoidea
Ancyloidea
(superfamily also
includes families
Physidae, Planorbidae)

Ancylidae

Subfamily

Genus

Ancylinae

Rhodacmaea

Ferrissinae

Ferrissia
Hebetancylus

Laevapecinae Laevapex

Banarescu's more conservative approach uses:

Superfamily Family

Subfamily

Tribe

genus

Acroloxidae

Acroloxus
Pseudancylastrum

Lymnaeidae Lancinae

Fisherola
Lanx, with subgenera

Lanx ss, and Walkerola

Lymnaeacea

Planorbinae

Ancylini

Ancylus
Ferrissia

Physastrini Pettancylus
Patelloplanorbis
Ancylastrum

Planorbidae Bulininae
Bulinini
Rhodacmeinae

Laevapex
Rhodacmaea

A few examples, all magnified:

Lanx subrotundata (Tryon, 1856) At 10 mm, relatively large for a freshwater limpet. Living
animal, (8x) and shells, 4x. From a river in Oregon flowing over bedrock. When in habitat, each
animal has a small tuft of algae fibers on the shell, though none was growing on other objects or
surfaces.

Ferrissia rivularis (Say,

1817)
Ferrissia tasmanica

Most of North America.

8x magnification.

Ancylus fluviatilis (Muller, 1774) 8x.

(Tenison Woods, 1876)

European lakes. Interior and side view,

apex on upper right.

New South Wales, Australia.

8x magnification.

Laevapex fuscus (Adams, 1841) Much of North

Ancylus sp. Southwestern Brazil, rivers. Right

America, lower elevations. Dark surface encrustations

are typical. 8x magnification.

image shows the inside surface of this

translucent shell. 8x magnification.

Pilsbry and Becquaert, (1927) use the following classification for African freshwater limpets.
They include 22 species in the genus Barnupia, endemic to Africa. Several of their nominal
genera were not included in Banarescu's accounting.

Ancylidae
........."Ancylus"
.....Ferrissiinae
........Ferrissia
........Kincaidilla
........Barnupia
.....Pseudancylinae
........Pseudancylus

Barnupia kimiloloensis Pilsbry

and Becquaert, 1927. Kimilolo
River, Africa. 6 mm. Courtesy
American Museum of Natural History.