GASTROPODA
SNAILS AND SLUGS
CLASSIFICATION- three subclasses
Class Prosobranchia
largest and most diverse
mostly marine; about 60,000 species
there are freshwater species in this order
(i.e., Pleuroceridae: Pleurocera, Campeloma, Viviparus, etc.)
Class Opisthobranchia:
exclusively marine, 11,000 species
Class Pulmonata
primarily freshwater and terrestrial, and marine limpets
the mantle cavity has been modified for air-breathing; have no gills
about 20,000 species
aquatic species have secondarily invaded water and must come to the surface periodically to breathe; no operculum
gill have been lost; the roof of mantle cavity is highly vascularized and acts as a functional lung
air gets in and out by the pneumostome
divided into 2 superorders:
Suborder Basommatophora
mainly freshwater species; some marine
one pair of tentacles with eyes at base of tentacles
Suborder Stylommatophora
terrestrial species
2 pairs of tentacles; eyes at end of posterior tentacles
some species have great reduction or loss of shell; these are the slugs
In both the Opistobranchs and pulmonates there is tendency for loss of larval forms, and a reduction of torsion; reduced to about 90 degrees
LOCOMOTION- there are 4 primary ways; First three used by freshwater species
Foot - movement by the foot; various wave-like peristaltic movements cause gastropods to creep along due to muscular contraction
Ciliary Action - cilia at bottom of foot; especially in small species that live on sand and mud bottoms
Operculum - those that have operculum use it by extending and anchoring it and foot contracts and pulls animal along; also used to dig
Swimming - by the use of extensions of the foot (fins) also called parapodia; best developed in opisthobranchs such as sea hares, sea butterflies
GENERAL MORPHOLOGY
Shell- structures and terms
1. shell orientation; types of spires
conical - as in limpets
planospiral - spiral along a horizontal plane; Helisoma
spiral - along a vertical plane; Pleurocera
2. Operculum - trap door to aperature
In Prosobranchia three types
concentric- rings in concentric circles; Viviparidae
paucispiral- rings few in number, usually a single loop; Pleuroceridae
multispiral- rings few in number, usually one, but many loops; Pleuroceridae
No operculum in Pulmonata
3. Columella - central "post" around which shell spirals
4. shell aperature orientation
dextral- opening on right side
sinestral- opening on left side; Physidae
5. Protoconch- "first shell"; shell that is laid down upon hatching, newly hatched snail is called a spat; first whorl
6. whorls
individual spirals of shell
body whorl- last whorl, where largest portion of body is located
morphology of whorls
rounded; Physa
carinate- with an edge, or two; Valvata
spines; Io
nodules; Lithasia
smooth; Campeloma
Umbilicus
perforate- with umbilicus
imperforate- without
Teeth- refers to structures along lip of body whorl of shell; not common in freshwater species- in some Planorbidae
6. Shell layers; laid down by mantle
Periostracum- outer horny layer, high concentration of organic matter
Prismatic- middle layer of calcium carbonate; molecules laid down vertically
Nacre- inner layer, lies against the body
NUTRITION- All gastropods have a radula and it is used for feeding; it is modified from one group to the next depending on the type of feeding that is involved
more primitive condition is that the radula is composed of several transverse rows of teeth which is used for rasping
in more advance gastropods, especially neogastropods, the radula is modified for predation
have radula consists of long sharp, barb-like teeth
In some cones radula developed into a harpoon-like structure used to inject venom into prey
gastropods are found in all parts of the food chain; freshwater gastropods tend to be herbivores or detritivores
Herbivores - found in both classes; snails are part of a small group of animals that produce cellulases (ie. enables them to digest cullulose)
feed on algae; rasping it off the substrate or may feed on the softer tissues of plants, using the radula to shred the food; mostly prosobranchs such as Pleuroceridae
some are able to feed on macrophytes, ie. Lymnaeidae; most graze (ie. scrape) on periphyton- especially green algae and diatoms
Carnivores - no freshwater species specifically carnivores; mostly marine species such as cones
Scavengers and Deposit Feeders- feed primarily on dead and decaying organisms or on detritus; found primarily in slow moving, silty habitats; primarily pulmonates, and some prosobranchs such as Camperloma
may have a proboscis that enables them to suck up detritus off the bottom
Suspension Feeders (Filter Feeders)- primarily feed on plankton; example is Bithynia tentaculata (Bithyniidae) that uses its ctenidium (gill) to filter phytoplankton from water; found in eutrophic lakes in New York
food is carried in by inhalant current over gills
gills secrete a sticky substance trapping food
in these the radula is usually very reduced; in a few species it seems to be lost
there is usually a well developed ciliary system in the gills and proboscis to transport trapped food to mouth
Effects of Plant Production by Snails
Gastropods are known to play an important role in plant, especially periphyton, production. Numerous experiments have shown a distinct difference in production in the presence vs the absence of snails
EXCRETION AND WATER BALANCE- Many archeogastropoda have 2 nephridia, but in all others the right one has disappeared or is very reduced and functions in reproduction only
because of torsion nephridia are located anteriorly in visceral mass
nephridia are sac-like structures that usually open into mantle cavity and wastes are removed by water currents
in pulmonates this is not possible because mantle cavity forms a lung so ureter is elongate and opens to front of mantle near the anus and pneumostome
freshwater species have to get rid of excess water so nephridia transport lots of water to outside
CIRCULATION
because of torsion heart is anterior in the visceral mass
in primitive archaeogastropods there are two auricles,
in all other gastropods the right auricle has become vestigial or lost (fig. 11-40, page 382 gives a diagram of typical circulation of blood in snail)
NERVOUS SYSTEM
in advance gastropods it is twisted (see Barnes Fig. 11-41, page 383) into a figure eight
as with other systems this twisting results in the loss or reduction of structures on the right side in this case it is affects the number of sense organs
sense organs include:
eyes- used to detect changes in light; not form images
tentacles
osphradia- used to detect food; located in mantle cavity near gills
statocysts- located near foot used for orientation
REPRODUCTION
Prosobranchia
most prosobranchs are dioecious with a single ovary or testis in visceral mass
males use the enlarged right tentacle as a copulatory organ, or may have a specialized penis (= verge) as in Valvatidae, Hydrobiidae, and Pomatiopsidae; Some have no special organ as in Pleuroceridae
Many lay egg masses; Pleuroceridae
Some are ovoviviparous such as Viviparidae- give birth to live young that developed in anterior fold of mantle
Some viviparids (ex. Campeloma) are parthenogenetic, which has adaptive advantages in certain environmental conditions; colonization of new habitats or in unpredictable habitats
The Valvatidae are hermaphroditic, but usually cross fertilization is the rule
Pulmonata
Pulmonates are hermaphrodites (monoecious), where copulation is the rule
sperm and eggs produced in an ovitestis that leave via an hermaphroditic duct (except in ancylids which have separate ducts; cross fertilization is required)
most are simultaneous hermaphrodites - produce eggs and sperm at the same time
eggs fertilized in hermaphroditic duct; internal fertilization
self fertilization
cross fertilization through copulation, most common when possible
eggs laid in gelatinous masses and attached to some substrate
Development:
in terrestrial and freshwater snails there is direct development with no larval stages
LIFE HISTORIES
Prosobranchia
either oviparous or ovoviviparous
some species are
annual- one year life cycle; reproduce once (semelparity)
perennial- may live 4-5 years and reproduce each year (iteroparity)
Pulmonata
oviparous hermaphrodites
typically annual (one year life cycle) and are semelparous, very few are perennial
typically reproduce in the spring and then die; ex. Lymnaea, Physa, Aplexa
some species have a single cohort, others may have two cohorts- one reproduces and dies in spring the other in the summer
DIVERSITY AND DISTRIBUTION (see table 10.1; page 291)
Prosobranchia
Pleuroceridae
widespread in Africa and Asia; greatest diversity in streams of SE U.S.
shell anatomy is used for genus level identification
operculum is corneus and paucispiral
Viviparidae
worldwide distribution
Campeloma (widespread), Lioplax (widespread), Tulotoma (restricted to Alabama) are endemic to North America
Viviparus is widespread in eastern North America, probably more common in North East
Ampullaridae
restricted to Florida, most common is kite snail, Pomacea
Valvatidae
11 species in North America
egg laying hermaphrodites
small shells, often carinated, dextral, multispiral operculum
Hydrobiidae
28 genera in North America; 152 species
dextral, paucispiral operculum, very small shells
Pomatiopsidae
one genus, Pomatiopsis, 6 species
amphibious, often found along shoreline
similar in appearance to hyrobiids
widespread
Pulmonata
Ancylidae
worldwide distribution
freshwater limpets, simple cone-shaped, sinestral shell
apex inclined to right
have modified gill (pseudobranch); secondarily derived; unique to this family in Pulmonates
Lymnaeidae
Worldwide distribution, most diverse pulmonate group
9 genera and 57 species
most have very thin shells, some get very large
mostly found in lentic waters
Physidae
worldwide distribution; ubiquitious in North America
Often referred to as having a single genus, Physa; in recent years it has been divided into 4 genera with 37 species
all are sinestral, most with very thin shells
Planorbidae
widespread
11 genera and 44 species
sometimes aligned with ancylids into a single family, Ancyloplanorbidae (See Branson reprint)
have hemoglobin
shells planospiral; range from 1mm (Neoplanorbis) to 30mm (Helisoma)
ECOLOGICAL CONSIDERATIONS
Limiting Factors
Calcium carbonate; best when alkalinity ranges from 20-600 mg/l; under 20 mg/l snails become stressed
pH- found in ranges of 6.7-9.0, most prefer 7.0-8.4; too low H+ ions will dissolve shell
Oxygen- not as much a problem with pulmonates since they rely on atmospheric oxygen; Prosobranchs- most need 5 mg/l or greater, few can survive periods as low as 2 mg/l
Temperature-closely associated with DO, usually not a problem in itself
Distribution
Major drainages with endemics
Great Lakes, St. Lawrence Seaway
Atlantic Slope
Gulf of Mexico; especially Coosa R. in Alabama
Mississippi; most species and genera here and in preceding drainage
Hudson Bay
Colorado R.
Interior Basin (Nevada, Arizona)
Columbia R
Pacific Slope
Artic Slope
Bering Sea
Important Mississippian Genera
Pleurocera
* Elimia (=Goniobasis)
Physa
Campeloma
Helisoma
Ferrissia
Habitat Preferences by Family
Two broad categories:
Important Lentic Families (most tend to have light shells, often very large, i.e. no current)
Planorbidae
Lymnaeidae
Physidae
Important Lotic Families (most have heavy shells used for ballasts; also tend to have high spires for the current)
Pleuroceridae
Viviparidae
Ancylidae