mollusks

molluscs

Mollusca
bltdjur

What is a mollusc?
Fundamental organization (hypothetical archimollusc): - shell secreted by a layer of tissue called the mantle - mouth and anus at opposite end (but in gastropods both anterior) - mantle cavity bears gills (but pulmonate gastropods have no gills) - above mantle cavity is the visceral mass with gut, nervous, circulatory and muscular system - shell is of calcium carbonite (calcite or aragonite) (but may be secondaryly lost) - shell typically external (but in some groups it became internal) - grow by accretion (calcium carbonate is added to the edge of the shell by the mantle) - generally marine (but also few freshwater terrestrial groups)

Mollusca

systematics

main groups

Mollusca

Bivalvia

Mollusca systematics Bivalvia

Bivalvia (=Pelecypoda, Lamellibranchia)

pair of valves (right and left valve) bilobed mantle valves articulate along a dorsal hinge line no head typically bilaterally symmetric (plane of symmetry passing between the valves, = commissural plane) prominent ventral foot known since the Early Cambrian, but diversify not prior to Ordovician but still not a very common faunal element during the Paleozoic

Mollusca systematics Bivalvia

Main features of the shell

muscle scars
homomyar heteromyar monomyar

ligament
internal external amphidetic prosodetic opisthodetic !

dentition
cardinalia lateralia

lunule
(escutcheon)

pallial line
sinupalliate integripalliate

beak
orthogyrate prosogyrate ! opisthogyrate

Mollusca systematics - Bivalvia

Basic for systematics are the gill type and the hinge dentition Gills:
Protobranchs (deposit feeders, most primitive) Filibranchs (suspension feeders) Eulammelibranchs (suspension feeders) Septibranchs (carnivores, most derived)

Mollusca systematics Bivalvia gill types

Dentition: Various types and subtypes taxodont dysodont isodont schizodont desmodont pachydont heterodont

Mollusca systematics Bivalvia dentition

Types of dentition
Taxodont many small similar teeth & sockets all along hinge plate (e.g., Glycimeris and Arca) Schizodont two or three thick teeth with prominent grooves (e.g., Trigonia) Dysodont small simple teeth near the edge of the valve (eg Mytilus) Heterodont few teeth varying in size and shape, distinquished as cardinal teeth, beneath the umbo, and lateral teeth which lie obliquely along the hinge plate (e.g., most recent bivalves) Isodont teeth very large and located on either side of a central ligament pit (e.g., Spondylus) Desmodont teeth very reduced or absent (e.g., Mya) with a large internal process (the chondrophore) carrying the ligament

Mollusca systematics Bivalvia dentition

taxodont
Taxodont many small similar teeth & sockets all along hinge plate (e.g., Glycimeris and Arca)

dysodont

Dysodont small simple teeth near the edge of the valve (eg Mytilus)

no teeth just crenulation

heterodont

Heterodont few teeth varying in size and shape, distinquished as cardinal teeth, beneath the umbo, and lateral teeth which lie obliquely along the hinge plate (e.g., most recent bivalves)

cardinalia and lateralia

isodont

Isodont teeth very large and located on either side of a central ligament pit (e.g., Spondylus)

two teeth correspond to two grooves

schizodont

Schizodont two or three thick teeth with prominent grooves (e.g., Trigonia)

teeth have crenulations (teeth with teeth)

desmodont

Desmodont teeth very reduced or absent (e.g., Mya) with a large internal process (the chondrophore) carrying the ligament

internal process (the chondrophore) carries the ligament

pachydont

Pachydont large, heavy and massive teeth (e.g., rudists)

Orientation of a bivalve shell

what is posterior anterior right left ? ligament typically posterior posterior adductor muscle scar stronger developed pallial sinus posterior / shell gaps posterior posterior part of shell typically better developed umbo (beak) typically points anterior (prosogyre) byssal notch anterior Oysters: left valve bigger/cemented

Mollusca systematics Bivalvia orientation

right

Mollusca systematics Bivalvia orientation

right

Mollusca systematics Bivalvia orientation

right

Mollusca systematics Bivalvia orientation

right

Mollusca systematics Bivalvia orientation

left

Mollusca systematics Bivalvia orientation

Ecology
marine and fresh water typically benthic, infaunal or epifaunal include burrowing, browsing, cemented, free lying, swimming, boring forms filter feeders, deposit feeders, carnivores

Mollusca systematics Bivalvia ecology

Infaunal bivalves
Both detrivorous and filtering strategies Most Palaeozoic groups are infaunal detrivores Probably the oldest of all bivalve life-modes Burry thorugh sediment with the muscular foot Extensions of the mantle (siphons) allow water transport Shell modified to specific substrate requirements

Infaunal bivalves
Water Venus is a shallow burrowing form with short retractable siphons. Sediment Mya arenaria is a sluggish bivalve which burrows quite deeply in firm sand or mud. Its long siphons can be retracted, but not all the way back into the shell

Water Sediment

Internal view of right valve

Shell features (deeper burrowers): Generally more elongate shells Some have gapes in the shell commissure to allow siphons to remain outside when shell is closed Dentition reduced

Shell Features (shallow burrowers): Equivalved Thick(ish) valves Adductor muscles roughly equal in size Commonly with strong external ornament

Note difference in size of pallial sinus between the two bivalves. (Generally the bigger the indentation the bigger the siphon and consequently the deeper the bivalve could burrow)

Internal view of left valve

Foot

Sessile Epibenthic bivalves

Attaches to hard subsrates and becomes immobile Many groups have evolved this lifemode independently Allows effective filterfeeding Mytilus (common blue mussel) and many others attach by chitinous threads (byssus) secreted by the foot Oysters attach by cementing one valve (left) to the substrate and adapt to the shape of the substrate

Motile epibenthic bivalves

Lie exposed on the seabed Mostly filterfeeders Acute sensory system including photophores (eyes) and sensory tentacles along the mantle edge Escape strategy: Rapid closure of the valves creates jetstream and the mussel can thus swim short distances Some Jurassic bivalves may have been permanent swimmers

Soft sediment recliners and mudstickers

Some byssally and cementing forms have evolved secondary soft sediment life-modes Larvae attaches to small objects and develops shapes that allows the bivalve to survive on the sediment surface

Gryphaea (devils toenail)

Pinnate bivalves

Reef-forming bivalves
Rudists (Jurassic-Cretaceous) reef builders Differential valves Cone-shaped right valve Left valve acts as a lid Probably had symbiotic algae like modern Tridacna Evolved from oysters?

Modern Tridacna clam

Reef-forming bivalves

Reef-forming bivalves

Rock boring bivalves

Several groups of bivalves can produce
livingchambers by boring through rock and wood Lithophaga calcareous substrates (corals, limestone etc.) Valves without gape Exclusively chemical excavation Pholadids All types of substrates Wood, corals, granite, lead cables, plastic, amber etc. Valves with wide anterior gape Excavation by movement (abrasion) Shell ornament of teeth and rockfragments wedged between them act as sandpaper

Cephalopoda

Mollusca systematics Cephalopoda

Cephalopods
most highly evolved molluscs (especially eyes and brain) a high level of cephalization (concentration of sensory and neural centers in the head) group includes the modern Nautilus, argonauts, squids, octopuses, cuttlefishes as well as the fossil ammonites and belemnites 2 main groups: Palcephalopoda (nautilids and endoceratids) Neocephalopoda (orthoceratids, ammonites, belemnites) typically bilaterally symmetrical shell, if developed, subdivided in chambers by septae chambers are connected by a tube (siphuncle) hyponome and tentacles are homologue to foot of bivalves and gastropods mouth with powerful horny beaklike jaws and a radula radula less developed than in gastropods since Late Cambrian

Mollusca systematics Cephalopoda

Neocephalopods

Octupus Spirula

Sepia (Cuttlefish) Loligo (Squid)

Shell remains

Palcephalopoda (Nautilus + fossils)

Shell terminology
shell wall peristome

aperture

growth line

septum protoconch

camera / chamber septal neck

p

on oc m ag r h

Mollusca systematics Cephalopoda shell morphology

liv in g

ch am be r

Mollusca systematics Cephalopoda morphology shell

The suture = junction between septa and shell wall

- most important for taxonomy and phylogeny of Ammonitoidea - particular types characterize distinct families and orders

saddles: point in apertural direction lobes: point backward prosuture primary suture

Mollusca systematics Cephalopoda morphology shell

Shape of shell

Mollusca systematics Cephalopoda

The cephalopod jaw

Modern Cephalopods have a horny beak, either two simple plates or more complex structures There is also a radula with rel. simple, undifferentiated teeth

Mollusca systematics Cephalopoda morphology shell

Classification

Old: Nautiloidea Ammonoidea Coleoidea Palcephalopoda (~Nautiloidea) Neocephalopoda (Orthoceratoidea, Ammonoidea, Coleoidea) Palcephalopoda shell well developed and large, originally slightly curved siphuncle was situated between the center and the ventral surface. siphuncle generally large with internal deposits (important tax. feature) Neocephalopoda siphuncle thin and empty phragmocone originally straight with the siphuncle situated at or near the center later the position of the siphuncle shifted to the ventral surface (Bactritida), the shell became coiled (Ammonoidea) the shell became internal, reduced or absent (Coleoidea)

Mollusca systematics Cephalopoda

Palcephalopoda (= Nautiloidea, + several Paleozoic groups, excl. orthoceratids)

Mollusca systematics Cephalopoda

Neocephalopoda (= Orthoceratoidea, Ammonoidea, Coleoidea)

Ammonoidea - Goniatitida

Mollusca systematics Cephalopoda

Neocephalopoda (= Orthoceratoidea, Ammonoidea, Coleoidea)

Ammonoidea - Ammonitida

Mollusca systematics Cephalopoda

Neocephalopoda (= Orthoceratoidea, Ammonoidea, Coleoidea)

Ammonoidea heteromorphic ammonites

Choristocerataceae
Late Jurassic

Spirocerataceae
Middle Jurassic

Ancyloceratina
latest Jurassic to end Cretaceous

Mollusca systematics Cephalopoda

Neocephalopoda (= Orthoceratoidea, Ammonoidea, Coleoidea)

Coleoidea

Coleoids have little skeletal material Consequently are rare as fossils Fossisl date back to the Carboniferous Probably derived from orthocone Neocephaolopds in the Devonian

Neocephalopoda (= Orthoceratoidea, Ammonoidea, Coleoidea)

Coleoidea Belemnitida

Belemnites were squid-like with internal shell (Phragmocone) The posterior of the phragmocone had mineralised deposits (rostrum or guard) The rostrum is a massive, calcareous structure and hence fossilise extremely well (contrary the phragmocone) Probably worked as counterbalance (compare darts) Belemnites were common in the Jurassic and Cretaceous No modern cephalopods produce a rostrum

Mollusca systematics Cephalopoda

Evolution

evolutionary explosion high diversity increase in size Plectronoceras

Mollusca systematics Cephalopoda evolution

Ecology
entirely marine active predators (all are carnivorous) active swimmers
swimming is by rapidly expelling water from the mantle cavity the water is forced out through the hyponome (jet propulsion)

Mollusca systematics Cephalopoda ecology

Swimming
swimming is by rapidly expelling water from the mantle cavity the water is forced out through the hyponome (jet propulsion)

Cephalopod eyes
Camera eye fully comparable to ours Famous case of convergent evolution Forms from skin in the embryo, ours from extension of the brain Nautilus has very primitive, pin-hole camera type eye

Homo sapiens

Octopus

Biostratigraphy
especially Ammonoidea and in the Mesozoic

Mollusca systematics Cephalopoda biostratigraphy

Polyplacophora

Mollusca systematics Polyplacophora

Polyplacophora (chitons)
primitive molluscs with eight, articulating (overlapping) aragonitic plates (except one Palaeozoic lineage had seven) generally oval in outline with a flattened body creeping foot, a primitive feature in molluscs radula, mineralized with magnetite head is poorly developed the girdle (perinotum), a band of muscular tissue, runs along the dorsal periphery embedded in the girdle are small calcareous spines, scales or spicules known since the Late Cambrian (isolated plates)

Mollusca systematics Polyplacophora

Multiplacophorans
Stem group polyplacophorans? Different numbers of sclerites Best know is Polysacos from the Carboniferous 17 plates

Polysacos

Polyplacophoran Ecology
marine, commonly occurring on rocks and seaweed in the intertidal zone few species have also been found at depths down to 5000 meters photoreceptor cells in the mantle and girdle. the animal is thus able to detect light, which it responds negatively to active at night, when they creep over rocks scraping algae and other microscopic organisms off the surface with their radula

Mollusca systematics Polyplacophora

Gastropoda

Mollusca systematics Gastropoda

Gastropoda
mollusks with a head and foot (the head-foot), and a mantle covering visceral mass head-foot can be withdrawn into the shell (sealed by operculum) typically with a univalve calcareous shell (maybe reduced, or pseudo-bivalved) shell generally coiled in some manner and external radula typically present Torsion is the single unique defining characteristic (synapomorphy) of the gastropods known since Late Cambrian

Mollusca systematics Gastropoda

Torsion
twisting of the body [it is entirely different from the spiraling of the shell
fossil evidence suggests that early, non-twisted molluscs already had coiled shells some modern gastropods have uncoiled shells, or even no shell at all]

all gastropods undergo torsion during some stage of their development - displacement of many interior organs - digestive tract became U-shaped (anus and nephridia moved anterior) - nervous system acquires a twisted appearance (streptoneury)

Mollusca systematics Gastropoda torsion

Torsion
Advantages: allowed the gills better access to water flow allowing the animal to withdraw more deeply into the shell the head was able to retract first (foot last, still able to swim) Disadvantages: anus and nephridia anterior the animal would be dumping its waste on its head

Mollusca systematics Gastropoda torsion

The radula
important taxonomic feature in modern gastropods no fossil radula confirmed, although there are descriptions composed of chitinous material and arranged as a long, coiled band consists of central, lateral, and marginal teeth

Mollusca systematics Gastropoda radula

Shell terminology
coiling: - dextral - sinistral

Mollusca systematics Gastropoda

Gastropod opercula

Traditional classification

Mollusca systematics Gastropoda systematics

Prosobranchia (shelled gastropods in which torsion is complete)

classification based on gill and radula types -- unfortunately! Archaeogastropoda: holostome aperture = no siphonal canal (since Cambrian)

Mesogastropoda: aperture typically with siphonal canal (since Ordovician)

Neogastropoda: aperture siphonostome, often very long siphonal canal (since Cretaceous)

Mollusca systematics Gastropoda classification

Modern classification
Incertae Sedis (primitive forms - Archaeogastropoda in part)
Order "Tropidodiscida" ("Bellerophontina" in part) Order Bellerophontida ("Bellerophontina" in part)

Subclass Eogastropoda (primitive forms - Prosobranchia / Archaeogastropoda in part)

Order "Platycerida" Order Patellogastropoda (Docoglossa) Order Cocculinida (polyphyletic?) Order Vetigastropoda

Subclass Orthogastropoda (all other gastropods)

Infraclass Neritimorpha (Archaeogastropoda in part) Infraclass Apogastropoda Superorder Heterobranchia Order Opisthobranchia Order Pulmonata Superorder Caenogastropoda(Prosobranchia in part) Order Architaenoglossa Order Neotaenioglossa Order Neogastropoda

Mollusca systematics Gastropoda systematics

Patellogastropoda

Neritimorpha

Cellana radians

Vetigastropoda

Amblychilepas scutella

Turbo (Dinassovica) imperialis Haliotis (Haliotis) midae

Caenogastropoda

Oliva (Oliva) sericea textilina

Pusionella vulpina

Morum (Oniscidia) exquisitum Turritella ungulina

Conus (Asprella) alabaster

Murex (Murex) aduncospinosus

Malea ringens

Heterobranchia Pulmonata Ophistobranchia

Glaucilla marginata

Philine angasi

Ecology
most are aquatic, marine, brackish and fresh water several groups lives on land (most are Pulmonates) marine forms typically live in shallow waters highest diversity in tropical waters but also known from arctic waters and hydrothermal vents in the deep sea one of the most adaptable forms with respect to: salinity preassure (water and air) temperature (water and air) humidity most are herbivores, but also carnivore (Muricidae, Naticidae, Conidae) and omnivore marine forms typically benthic, but also free swimming and floating forms freshwater and terrestrial forms at least since Carboniferous

Mollusca systematics Gastropoda ecology

Palaeozoic gastropods

Relatively rare Shell usually structurally weak: With selenizone or anal slit Lacking columella (central strengthening rod connecting whorls) Platyceratids Large, loosely coiled shell Uneven margins Life attached to crinoids

Palaeozoic gastropods

Bellerophontids Planispiral coiling Selenizone and deep sinus Selenizone often raised Extinct Name derived from ancient Greek hero Bellerophon in recognition of the similarity to a greek helmet

Bucanella nana

Sinuites

Bellerophon

Modern gastropods

Mollusca systematics Gastropoda

Cone shells

Hunt with poisonous harpoons Poison sometimes extremely potent (deadly to humans) Prey is ingested whole or scraped with radula

Patellids

Cap-shaped shell Sticking to rocks and other hard things Foot modified to function as a sucker Why? Protection Conserve moisture Feeding by scraping algae Secondarily untorted Obs! Convergent evolution

Predation by Gastropods
Several groups of gastropods feed by drilling holes in mollusc shells Muricids are epibenthic with often highly ornate shells. Drill holes with straight sides Naticids are infaunal with very smooth, rounded shells. Drill countersunk holes by combining acid with radular activity Naticid

Muricid

Mesozoic marine revolution

Predator-prey arms race Jurassic to present Evolution of new predators (e.g. tools) - Crab and lobster claws Today: More shells are damaged than in Palaeozoic Led to new mollusc adaptions - glossy shells - varices on aperture - narrow aperture New inventions forced the opponent to develop new counter methods Affected all benthic marine animals