molluscs peter shaw. phylum mollusca n one of the most important phyla in terms of species richness,...
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MolluscsPeter Shaw
Phylum Mollusca One of the most important phyla in
terms of species richness, and nearly ubiquitous in most ecosystems
They invariably have a eponymously soft, mucus-covered body. Many forms have a hard, calcareous shell protecting most of the body.
The mollusc body Is unsegmented Coelomate, but coelom is reduced to
pericardium + gonadial cavity. The main body cavity is a haemocoel.
Primitively: shelled, with 2 broad body regions:
1: a non-muscular visceral hump staying protected inside the shell, wherein most action is ciliary.2: a muscular foot, with mouth + sense organs at anterior end which protrudes from the shell
The ancestral mollusc A favourite parlour game for invert.
Zoologists is to create a hypothetical ancestral mollusc.
In 1957 a monoplacophoran Neopilina galatheae was dredged up from 5000m off Mexico, and was found to fit the bill - except that, rather embarrassingly, it showed signs of metamerism.
Ancestral mollusc (hypothetical)
Phyletic affinities
Are very much disputed, but current best guess is close to some obscure worm-like phyla: sipunculids, echiurids, and more distantly annelids. These are now classed with lophophorate phyla as a subgroup within the lophotrochozoa.
This scenario assumes a ciliary/gliding animal which in one radiation acquires a shell, the other evolves lophophorate filter feeding
Respiration
Most molluscs are aquatic and bear paired gills or ctenidia, lying in the mantle cavity. These bear many triangular leaflets, and water is carried over these by ciliary action.
Water is “tasted” before flowing over the gill by a sense organ the osphradium.
The anus and renal pores lie in the exhalant stream.A single ctenidium
(typically triangular, leaf-like)
Respiration contd.
Neopilina has no ctenidia, but 5 pairs of muscular gills along its body. Primitive or derived?
Terrestrial gastropods often lose ctenidia, and have “lungs” in their place.
(And a few of the air-breathing pulmonates then returned to the sea!!)
Pneumatostome, leadingTo mantle-cavity ‘lung’ in banana slug
Mouthparts Usually consist of a unique molluscan organ: the radula. This is a muscular cord which bears sharp chitinous teeth, which rasps slowly away at the surface of food grating fragments into the mouth.
The radula is born on a supporting structure the odontophore.
More on mouthparts
Radula teeth vary immensely with diet, an extreme being Conus, whose teeth are hollow venom-injecting needles.
In Chitons, the radulae are tipped with magnetite for extra hardness.
In cephalopods the radula is still present but supplimented by a parrot-like beak.
Feeding
Primitively this involved food particles being scraped off a surface by the radula/odontophore and carried in a mucus rope into the stomach, where it is size-sorted by cilia.
Filter-feeders retain the mucus rope, but collect particles off the ctenidia.
Cephalopods bite lumps of flesh with a chitinous beak prior to radula.
The shell This is invariably calcium carbonate (aragonite, not
calcite - the crystal structure differs slightly) reinforced with chitin.
The superficial appearance differs hugely between groups, from a tight spiral through loose tubes to nearly flat.
In fact one growth equation with 3 biologically realistic parameters can fit the whole spectrum
One weird footnote from a weird system: a snail, unnamed in 2003, inhabiting the deep-sea black smokers, has been found to supplement its normal carbonate shell with scales of iron sulphide.
Classification This seems to have changed over
the years, due to increased ‘splitting‘ of obscure minor forms in recent years
To a good first approximation there are 5 main classes, but you need to know of another three oddball classes.
Principal mollusc classes
C e ph a lo po dsO c top i, squ id s, cu tt le f ish e tc
G a stro po dsS lu gs , sn a i ls w ink le s e tc
P o lyp lacop ho raC h ito ns , co a t-o f-m a i l she lls
S cap ho po dse le ph an t tu sk she lls
L am m ellib ran ch iab iv a lv e s: m u sse ls , o yste rs e tc
Mollusca
Minor classes
M on op lacop ho raN eop i lina
n e om e n io rp hs C h ae to d erm o m o rp ha
A p la cop ho ra
M o llu scs
Polyplacophora
Chitons, or coat of mail shells Primitive marine molluscs with 8 small
dorsal shells, partly or wholly buried in flesh.
This is not segmentation but an adaptation to curling in tight crevices
Feeding is by grazing on algae. Mantle cavity runs along side, almost up
to head, with paired gills along its length
Scaphopods
“Tusk shells”: 350 spp mantle secretes a tubular shell,
open at both ends Burrow in soft marine sediments,
collecting food particles with sticky clubbed captactula
Bivalvia = Lammellibranchia = Pelecypoda C. 20,000 spp Sedentary/ sessile molluscs encased in
paired shells Lacking eyes, radula and tentacles,
though these may re-evolve around mantle edge
Foot retained, often used to burrow into sediment
Live by filter feeding, mainly using ctenidia as filtering surfaces
Gastropods
The most numerous group of molluscs, with c. 76,000 species (75%)
Mainly shelled, except for the nudibranchs
The shell is always single, usually spiral
Locomotion is by muscular waves passing along the foot, except for a few tiny spp that use cilia
Gastropods, 2
A typical gastropod is a squat mollusc crawling with its muscular foot.
The cephalic sensory organs and radula are well developed but can retract fully into the shell
Eg common limpet Patella Many gastropods have a
calcareous/chitinous plug to the shell - the operculum
Torsion All have undergone a
curious modification of body form known as torsion
The visceral hump is rotated 180o so that the mantle cavity, gills and anus all point anteriorly
Perhaps to allow the head to retract into the generous space of the mantle cavity + allow osphradium to sense oncoming water
Gastropods: torsion
But having your anus discharging over your head is poor design, even for a mollusc
The solution is to modify the exhalent stream: evolve a special slit in the shell lose one ctenidium (R side) giving a
unidirectional flow Loss of R gill => loss of R kidney, osphradium
+ R side heart L (inhalent) mantle protrudes as a siphon
Many spp have undergone 90o detorsion!
Gastropod orders
P roso bra nchsa n ce s tra l ga stro po ds
O p isth ob ran chsM ain ly m a rine
p u lm on atesm ain ly te rres tria l
G a stro po ds
Prosobranchs
Mainly marine “snails”, with traditional body/shell design
Most graze algae, sessile animals, or sediment
Many have notable shells, ie cowries
One order Stenoglossa have effective predators: Conus spp hunt+kill fish!
Opisthobranchs
Marine, shell-less: sea slugs, sea hares, pteropods etc.
Some are pelagic, hunting cnidaria one group are pelagic filter feeders
using mucus nets Ctenidia are reduced or absent:
the body surface is often adequate, or 2ndry gills have evolved
Pulmonates
Mainly terrestrial radiations Mantle cavity becomes an air-
breathing lung with a contractile opening the pneumatostome
Most have a shell, but stylommatophora have lost it: land slugs
A few pulmonates have returned to water!
Class: Cephalopoda NautilidaNautilus
Am m onitesextinct
Octopoda
Teuthidasquids
Vam pyrorpha'Vam pire squids'
Sepiidacuttlefish
Cephalopods
The most intelligent, fastest moving and highly modified molluscs
all are marine predators with good/ excellent vision
include the largest invertebrate (giant squids)
foot is modified into multiple tentacles with suckers
swim by jet propulsion
Cephalopods We are lucky to have alive an
ancestral form, Nautilus (6 spp). This has a spiral shell used for protection and buoyancy.
The eye is a pinhole camera with no lens but a contractile iris
It lives as a deep water scavenger, probably explaining its survival of the K/T boundary eventIts fossil ancestors had straight shells (<= 4m long)
Ammonites
Were large open-water Nautiloids that went extinct at the end of the Cretaceous
Up to 2m diameter shells which differed from Nautilus by heavy sculpturing of sutures
We can only guess at their ecology - ?pelagic hunters
Some big bulky forms ?plankton feeders Evolutionary trends of their shells are
complex if not haphazard!
Coleoidea
Is the sub-class containing the remaining cephalopods octopoda teuthida sepiida Vampyromorpha These have a closed blood circulation,
non-ciliated gills, a good compound eye and well developed CNS. Are oddly semelparous
Cephalopod sex
These animals have some remarkable reproductive behaviour.
Sexes are separate and fertilisation is internal, but there is no penis. Instead one of the male’s arms is differentiated (usually with smaller suckers at the tip), is used to pull the torpedo-shaped chitinous sperm packet out of his genital opening, and insert it into the female’s body cavity.
In some deep sea squids the sperm packet may be jammed into non-standard regions - the mantle, or a tentacle base. How this effects sperm transfer is still unclear - these animals are effectively impossible to study alive.
Note the reduced suckers on the hectocotylus arm
Hectoctylus arm onSouth African diamond squid