publication 420-530

Sustaining Americas Aquatic Biodiversity

Freshwater Snail
Biodiversity and Conservation

Paul D. Johnson, Research scientist, Tennessee Aquarium Research Institute, Cohutta, Ga.

ix hundred fifty different species of snails are

widely distributed across the streams, rivers,
and lakes of North America. There are unique
species associated with every type of aquatic habitat from the Canadian Arctic to the Everglades of
Florida. However, the greatest species richness is
associated with flowing waters (streams and rivers). The
number of North American
freshwater gastropods (one or
no-shell mollusks) is the richest
in the world, rivaled only by the
river systems of Southeast Asia.
North American freshwater snails
represent a rich natural heritage of
global importance. Freshwater snails
are also an important food source for
many fish, turtles, and other species of
wildlife. Finally, because of their sensitivity to certain chemicals, many species are
excellent water-quality indicators.
Unfortunately, freshwater snails like many other river
species across North America are threatened. The
abundance and variety of snails have dramatically declined over the last 80 years, especially those species
that inhabit streams and rivers. Approximately 60 species of freshwater snails are presumed extinct, 20 are
on the federal endangered or threatened species list,
and another 290 species are of conservation concern
nationwide. In other words, 9 percent of all freshwa-

ter snails are extinct and 48 percent are conservation

targets. This rate of imperilment exceeds every other
major animal group in North America, even freshwater mussels of which 42 percent of all species are
conservation targets.
The decline of freshwater snails began in the early
twentieth century. Dam construction and other
channel modifications, siltation, and industrial
and agricultural pollution have all degraded the
river habitats on which most species depend.
As a result, the species richness and the abundance of freshwater snails have declined
dramatically. This decline has been especially pronounced in the Southeastern
United States where these animals are
most diverse. The continued loss and
decline of freshwater snails and other
river species (mussels, fish, crayfish)
are testament to the fact that despite
significant water-quality improvements
made in the last 25 years, much work
remains if we are to halt species losses.
Conservation and recovery efforts for freshwater snails include artificial culture, water pollution
control, and most importantly, habitat protection and
restoration. Cleaning waterways not only improves
the habitat for snails and other aquatic life, but it also
improves the quality and supply of water for human
consumption. Your help is required to assist conserva-

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tion and recovery efforts aimed at rivers and streams,

and the animals that inhabit them. Adopt a stream,
become involved with a local river conservation
group, or assist your state natural resource agency
with water-quality measurements.

Freshwater snails come in a variety of shapes and

sizes. Some snails are almost completely round in
shape, but most species are generally conical. Adults
of some species are smaller than the head of a pin and
others larger than a baseball.

What Is a Snail?

Along with size, shape, and coloring, structural

shapes such as large tubercles, spiny projections,
folds, sharp ridges, and fine lines on their elaborately
sculptured shells are what give some individual species their distinctive forms. These ornate shell structures are formed while the snail is building its shell.
They add strength to the shell to resist crushing by
predators (i.e., fishes and crayfishes). In contrast, the
pearly inside of the shell (nacre) is very smooth and is
usually a translucent white to light pink or even blue.
The interiors smoothness cushions the hard shell
against the snails soft body. Many species have internal striping on the nacre, which usually is also visible
on the outside of the shell.

Freshwater snails belong to

a larger group of
shelled animals
called mollusks.
Together, freshwater,
saltwater, and terrestrial
snails number well over 50,000
individual species, by far the largest group of
mollusks. Snails are soft-bodied animals that carry
with them a protective shell. This shell is composed
primarily of calcium and other minerals the snail has
eaten and hardened into a well-formed shell. The
soft-bodied animal is permanently connected to the
shell by a strong ligament and other soft tissues. The
snails shell is used for protection from environmental threats and predators.

The snails soft body is divided into three distinct

sections. A well-defined head is connected to a large
muscular foot. The foot is the most visible external
feature of the snails body and is the animals mode of
locomotion. A snail propels itself with this single foot,
which is controlled by hydrostatic action inside the
snails body. Additionally, the foot surface is covered
with tiny projections (cilia) that assist the gliding motion. Snails feed along the bottom as they move, partially clearing a path for the foot. As the foot passes
over the substrate, it leaves a trail behind the snail.
These trails are commonly visible in soft sediments or
across hard surfaces such as rocks or submerged trees.

Most species of freshwater snails originally evolved

from saltwater habitats, although several species
evolved from terrestrial habitats to freshwater. Snails
usually play a dominant role in the ecology of freshwaters by providing food for many other animals
and by grazing on vast amounts of algae and detritus
(debris). They are critical to normal ecological processes in rivers. Their unusual common names such
as: banded mysterysnail, applesnail, pagoda slitsnail,
knob mudailia, rough hornsnail, interrupted rocksnail,
among many others often belie their importance as a
food source to other aquatic animals and as indicators
of water quality.

The third distinctive section, the internal organs, is

concealed inside the shell. The internal organs of a
snail include a heart for circulating blood; a complete digestive system with a distinct mouth, stomach,
and intestine; a reproductive system; and a gill or
other respiratory surface used for oxygen exchange.
When a snail is threatened by a predator, the head
is retracted into the shell first, followed by the foot;
thus the entire body can retreat inside the shell. Some
snails even have a hard lid on their tails called an
operculum (oh-pr-ky-lum) that is made of protein.
The hard operculum covers most soft tissue of the
foot exposed to a predator when the snail has retracted into its shell.

Structure and appearance

Freshwater snails generally are dull colored, especially in comparison to their marine and terrestrial
cousins. Their color generally varies from yellowbrown to black, but some species have stripes and
other distinctive markings.
The shell is composed primarily of calcium carbonate (similar to chalk) and other minerals extracted
from the snails diet. The shell is coated with a horny
epidermal (outer) covering called the periostracum
(pair-i-ahs-truh-kum). This covering is what gives
the shell its color.

rivers of Southeast Asia appear to contain nearly as

many species. Additionally, new species of freshwater
snails are discovered each year. In the United States,
the abundance of species diversity lies in the river
systems of the Southeast.

Freshwater snails with an operculum are descended

from marine ancestors and extract oxygen from the
water with a single gill. These operculate snails are
the most diverse group, comprising about two-thirds
of all North American freshwater snails. They have
separate sexes and a short reproductive season. In
general, these species are slow-growing, longer-lived,
and very sensitive to environmental changes. These
species are distinctly distributed about separate river
systems and most major drainages have their own
unique species.

The Mobile River Basin

The rivers and streams of the Mobile River Basin
(streams in Alabama, Georgia, Eastern Mississippi,
and Southeast Tennessee) contain the greatest species
richness of freshwater snails in North America. The
Alabama, Black Warrior, Coosa, Cahaba, and Tombigbee river drainages together comprise
the Mobile River Basin. Although the
Mobile River Basin is about 3.5
percent the size of the Mississippi River basin, it contains 120
different species of snails. In other
words, over 18 percent of the species found in all of
North America are located in a basin with only about
1.5 percent of the nations surface area.
Unfortunately, some 37 species of snails in the basin
are now thought to be extinct. The construction of
several large hydroelectric dams on the Coosa River
were responsible for driving most of these species to
extinction. In addition, seven species of freshwater
mussels have been lost from the Mobile River Basin
in the past 80 years. The loss of 44 species of freshwater mollusks from the Coosa River Basin is widely
considered to be the single largest recent extinction
event in the United States.

In contrast, snails that lack an operculum evolved

from terrestrial ancestors and breathe with a modified
lung. These species are hermaphroditic, that is each
snail possesses both male and female reproductive
organs, and have an extended breeding season. They
are fast-growing, short-lived, and often less sensitive
to their environment. Unlike operculate snails, these
species are generally not limited by river drainages,
and most species are found across distinct drainage
basins.

Distribution and Diversity

Freshwater snails can be found living at the bottoms

of our largest lakes and rivers as well as the smallest
streams and ponds. Some species have been recorded
at depths of over 100 feet. The over 650 species in
North America are found from deep, interior Canada
near the Arctic Circle to the subtropical regions of
South Florida and Mexico. North America is the
Earths center of freshwater snail diversity. Only the
Spire

Egg Clutch
Tentacle
Head

Radula

Foot

Main Body Whorl

Figure 1. Major anatomical features of the spiny river snail (Io fluvialis).
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However, at specific localities in the Mobile River

Basin, snail diversity can exceed 12 different species.
Species diversity at these selected sites in the basin
is probably greater than at any other location in the
country. Additionally, snail densities can exceed more
than 50 per square foot of river bottom at certain locations. Yet, even this density can be exceeded at certain
locations within the Ohio River Basin.

and firm river bottoms, some prefer the soft substrates

more common to ponds and lakes. A few wideranging snail species can easily survive in polluted
habitats.
Snail species richness is greatest in large- to mediumsize rivers. Within the Tennessee and Mobile River
basins, it is not unusual for 10 or more species to be
present at a single location. Locations with stable,
clean river bottoms, clean water, high oxygen concentrations, and hard water usually support greater
species richness and densities. Clean, clear water, free
of sediments, is important to maintaining healthy snail
populations. Snails can live in rivers at depths of more
than 40 feet, as long as the water is clean enough to
allow light penetration to that depth. Generally, only a
few isolated locations in the Tennessee River contain
snails at depths greater than 20 feet, because of the
excessive nutrients and sediment.

The Ohio River Basin

The rivers and streams of the Ohio River Basin are
located across Northern Alabama, Illinois, Indiana,
Georgia, Kentucky, Mississippi, New York, North
Carolina, Ohio, Pennsylvania, Tennessee, and Virginia. These rivers and streams are home to the second
most diverse assemblage of freshwater gastropods
in North America. The Ohio River Basin supports
over 75 species of freshwater gastropods, or over 12
percent of all North American species. This includes
about five species that are now considered to be extinct. Once widespread and with populations numbering in the billions, many remaining river species
are limited to small ranges. In fact, range reductions
greater than 80 percent are common for many operculate species.

How Do They Reproduce?

Because freshwater snails occur across a variety of
freshwater habitats, they have adapted different reproductive strategies. There are basically three different
modes of reproduction commonly used by freshwater
snails.

The major rivers of the Ohio River Basin include the

Allegheny, Big Sandy, Cumberland, Green, Kanawha,
Kentucky, Monongalia, Muskingum, Tennessee, and
Wabash, which drain portions of many states. The
Tennessee Basin alone drains streams from seven
states (Alabama, Georgia, Kentucky, Mississippi,
North Carolina, Tennessee, and Virginia) and contains
the largest species richness of any Ohio River Basin
tributary. Species richness in several tributaries of the
Tennessee River is exceptionally high. These include
Limestone Creek in North Central Alabama, which
contains 22 species, including two federally listed
species, and the Duck River Basin of Central Tennessee, which contains 23 species. Snail densities in
these systems can easily exceed 100 individuals per
square foot of river bottom.

Most species of
snails are separately sexed,
and males
fertilize the female through direct copulation. Females
attach their eggs directly to firm, clean substrates such
as a rock, log, or aquatic plant, usually in shallow
water. Egg clutches are commonly laid on the shells
of other individuals. Egg clutches can contain two
to over 300 eggs, depending on the species. Females
can lay eggs in pairs, in a circular concentric pattern,
or even in a large line several eggs across and several inches in length. A female snail can take several
hours to deposit eggs for a single clutch.
For species that lay discrete clutches, the eggs are
deposited over a period of one to three months, usually in the late winter and early spring. Larger, older
females tend to lay more eggs per clutch. Warmer
water temperature plays an important role in egg
production, but day length also may be an important
factor. Individual eggs are small, a little larger than
1/75 inch in diameter. The eggs may take one to five
weeks to hatch depending on water temperatures. The

Where Are They Found?

Freshwater snails occur across a variety of habitats,
reflecting the wide-ranging biology of many different species. Many species spend their entire lives in a
few square meters of habitat, making them extremely
vulnerable to localized environmental habitat degradation. Although most species prefer clean, stable,
4

debris, root mats, and submerged plants. The ingested

algae and organic matter is mixed with a steady
stream of mucus as it enters the mouth. As their feet
glide most easily across firm surfaces, the snails feed
directly on top of the surface on which they are crawling.

warmer the water, the faster the eggs will hatch. A

juvenile snail is about 1/100 inch at hatching, but rapidly grows its first year to several hundred times its
birth size. Rapid juvenile growth in shell size to over
1/4 inch wide in the first year is not uncommon.
Another reproductive strategy of some operculate
species is for females to birth live young. In reality,
an egg hatches inside the female and the juvenile
snail grazes in a special pouch inside the mothers
body. After a few weeks of feeding, a juvenile snail
about 1/4 inch in length crawls outside the mothers
body. Female snails can produce several eggs simultaneously, and up to a dozen small juvenile snails
can be inside the female during the summer and fall
months. Like other operculate snails, these species
have separate sexes and the male uses a modified
tentacle as a penis. Juvenile snails emerge from the
female fully functional and ready to feed.

Many snails are adapted to move and feed in soft

sediments. They generally feed on tiny bits of organic
debris that are imbedded in fine sediments. Snails
will ingest sediment and organic material imbedded in a strand of mucus. This mucus stream flows
directly into the mouth. In rivers with heavy amounts
of suspended organic matter, snails may feed by lying
on their shells and turning their foot up into the water
column. Food collected on the bottom of a snails
foot is moved into the mouth through a strand of mucus. In this manner, snails can be suspension feeders,
directly collecting food from the water column.

Other species of freshwater snails have yet another

mechanism for reproduction. For these species, each
snail contains both male and female reproductive
systems, and all individuals can lay eggs. However,
individual snails do not self-fertilize, and sperm is
exchanged between individuals. Eggs are laid in
large, clear gelatinous clutches that can easily exceed
1/4 inch in diameter. Individual eggs and juveniles are
generally larger than those of other snails at hatching.
The total number of eggs is dependent on clutch size,
but the clutches of some species contain over 50 individual eggs. As long as the food is abundant, these
species generally produce eggs continuously from the
late spring into the early fall. Not surprisingly, these
species are widespread, commonly encountered, and
rarely of conservation importance. However, these
snails are a very important and abundant food for
many crayfishes and fishes.

How Long Do They Live?

Freshwater snails can vary wildly in their longevity, which is dependent on the type of snail and the
habitat in which it occurs. In general, snails that
possess an operculum live longer than species without one. The maximum age of many species living
in streams and rivers can exceed five years, whereas
most species that prefer ponds and lakes live less
than one and a half years. As a general rule, the more
nutrients available to the snail, the faster it grows, and
the shorter its lifespan. Snails living in nutrient-poor
habitats on average tend to live longer.
Determining the age of a freshwater snail is difficult,
and information about the maximum age for most
species is lacking. River species usually live from
two to six years. Io, the spiny river snail (see figure
1) may live well beyond 15 years, and is likely the
longest-lived of all North American freshwater snails.

What Do They Eat?

What Good Are They?

Whether they live in lakes, ponds, rivers, or streams,

most snail species eat small bits of algae and organic
debris found on the bottom and sides of the river
channel or lake. Freshwater snails actively graze a
variety of surfaces, scraping their food with a specially modified tongue called a radula. The radula
is covered with tiny, hard, blade-like projections that
scrape the surface. These tiny blades tear small bits of
algae and organic material from these surfaces. Most
snails graze on firm substrates, such as rocks, woody

The value of freshwater snails lies in their ecological

role in the aquatic food web and in maintaining water
quality. In healthy river ecosystems throughout the
Southeast, freshwater snails number in the millions
and serve as an important source of food for other
wildlife. Through sheer numbers, they contribute
greatly to the nutrient exchange processes in these
systems. Primarily by controlling algae growth, they
also maintain water quality and clean substrates for
5

other bottom-dwelling animals such as freshwater

insects.

shoal habitats on which snails depend, and it can also

increase water velocity, which can dislodge snails
during flood events.

Snails are a common food item for many species

of ducks, fishes, crayfishes, and turtles. In fact, one
group of threatened turtles that lives in the rivers
draining to the Gulf of Mexico relies heavily on snails
as food. Freshwater snails also are the major food
item for the snail darter, a small, endangered fish in
the Tennessee River system. Whether they are eaten
by turtles, fish, or dozens of other wildlife species,
freshwater snails play an important role in the aquatic
food web.

Dredging refers to the physical removal of natural

stream and river bottoms. Dredging removes gravel
or sand in attempts to control localized flooding, or to
deepen a river channel for boat navigation. Dredging
physically removes prime shoal habitat and destabilizes upstream riverbanks. In other words, when you
dig-a-hole in a river or stream, there is a natural
tendency for that hole to fill with materials stripped
from upstream banks. As a result, it is very common
to see collapsed stream and riverbanks, with dozens
of trees falling into the channel, upstream of recent
dredging activity. Riverbank instability is most pervasive in small- and medium-size rivers, because larger
rivers are more stable.

Snail Killers
The rapid decline in diversity and abundance of freshwater snails is attributed to a variety of human degradations of their natural habitats. The most prominent
impacts include: (1) dams and impoundments, (2)
channelization and dredging, (3) sedimentation and
channel instability, and (4) water pollution.

Many states consider stream and river bottoms to be

state property, and it is illegal for anyone to build a
dam or otherwise modify a stream or river channel
without permits. Any damming, diverting, channel modification, wetland draining, gravel removal,
or other direct habitat impact to a stream or river
requires permits from the U.S. Army Corps of Engineers and other federal and state agencies.

Dams and impoundments

Dams turn rivers into reservoirs and modify aquatic
habitat beyond the tolerance level of most river snails.
Eliminating current flow and increasing water depth
causes fine particles to settle on the reservoir bottom. Heavy layers of fine sediments literally smother
snails. Floating algae and other suspended materials
limit sunlight penetration to the lake bottom, reduce
algae growth on bottom substrates, and eliminate
the snails primary food source. In deep reservoirs,
the waters become layered with colder, oxygenpoor waters on the lake bottom, further reducing
any chance of snail growth and reproduction. Many
river species require flowing waters before females
will even deposit their eggs. Dams also are effective barriers for snails moving up-river and fragment
snail populations into small, isolated groups that are
more vulnerable to adverse impacts. Dams dont just
eliminate habitat for snails, they negatively impact
river species of mussels, fishes, crayfishes, turtles,
and salamanders.

Sedimentation and channel instability

As trees and other vegetation buffers are removed
from stream and riverbanks, the bare sediments exposed become vulnerable to removal during flooding
events. These exposed banks often wash away in
large chunks, depositing sediments in the river. If
left uncorrected, the stream bottom next to an eroding bank also becomes very unstable, and readily
moves during even minor flooding events. Shifting stream bottoms dislodge and bury adult snails,
juveniles, and eggs and reduce snail habitat. Streams
and rivers with high sediment loads are literally buried under layers of soft sediments and are devoid of
aquatic species that require firm substrates to move,
feed, and breathe.

Water pollution

Channelization and dredging

The specific effects of toxic

compounds
on freshwater snails

Channelization refers to the straightening of rivers

and streams by removing the natural bends and meanders of a normal channel. This straightening process
usually eliminates habitat for all but the hardiest of
snail species. Channelization destroys the shallow6

are poorly understood. However, the impacts of petroleum, certain metals, and agricultural fertilizers and
pesticides on freshwater snails are well documented
and can be lethal to snails in very small amounts.
Frequently these toxic compounds are attached to soil
sediments that are washed into streams during storm
events.

residential, urbanization, highway, mining, irrigation,

industrial development, etc.) if not properly managed, can have profound impacts on local rivers and
streams. Please report any suspected water pollution
problems immediately to your state authorities. Any
one of the following conditions could indicate a waterquality problem with your local stream or river.

What Can You Do?

1.
2.
3.
4.
5.
6.
7.
8.

Public involvement is critical to the protection of the

aquatic systems on which freshwater snails and many
other aquatic animals rely. Working to clean up and
restore stream and river habitats has many benefits
beyond wildlife conservation. Freshwater is a vital and
limited resource, essential for all kinds of life, including our own. Less than one percent of all water on the
planet is freshwater, so the degradation of our rivers
and streams limits our own water supply. Polluted
waters, which are still prevalent in some areas of the
United States, result in increased water treatment costs
to the taxpayer. Therefore, it is in everyones interest
to protect our aquatic resources and our water supply.
Get involved with your local watershed conservation
group. For information on the watershed group nearest you, please contact:

Muddy or unusually cloudy water

Petroleum on the water surface
Large numbers of dead fish
Excessive algae growth
Unusual or pungent odors
Loss of water volume
Brown foam on the water surface
Large amounts of litter

By helping to keep rivers and streams clean, you will

not only improve your local water quality, but you will
also improve habitat for all kinds of freshwater wildlife, including snails. The future of our rivers depends
on it. Remember nearly everyone lives downstream.
For more information about freshwater snails on the
Internet, please visit any of the following websites:
Freshwater Mollusk Conservation Society
http://ellipse.inhs.uiuc.edu/FMCS/index.html

The River Network

4000 Albemarle Street NW, Suite 303
Washington, DC 20016
(205) 364-2550

Freshwater Gastropods of North America

http://www.cofc.edu/%7Edillonr/fwgnahome.htm

Or

Freshwater Mollusk Bibliographic Database

http://ellipse.inhs.uiuc.edu:591/mollusk/biblio.html

The Nature Conservancy

Freshwater Initiative
6114 Fayetteville Road, Suite 109
Durham, NC 27713
(919) 484-7857

Acknowledgements
We greatly appreciate the editorial reviews of Adam
K. Downing and Nancy Templeman, Virginia Cooperative Extension, and the support of Randy Rutan
and Hilary Chapman, National Conservation Training
Center, U.S. Fish and Wildlife Service.

Because many local and state water monitoring agencies are understaffed and overwhelmed, they need
help monitoring rivers and streams. You should also
be aware of changing land-use and water practices
in your watershed. Land-use changes (agricultural,

Art illustrations by Sally Bensusen, Mark Chorba, and

Karen J. Couch.

Reviewed by Michelle Davis, research associate, Fisheries and Wildlife

7