Hickman, Miller

AmirV
Contents
Porifera ................................................................................................................................................................................................ 3

REPRODUCTION .............................................................................................................................................................3
Asexual: ................................................................................................................................................................3
Sexual: ..................................................................................................................................................................3
Larva types: ..........................................................................................................................................................3
GENERAL ASPECTS ..........................................................................................................................................................3
SKELETAL FRAMEWORK ...................................................................................................................................................3
CELL TYPES....................................................................................................................................................................4
Amoebocytes: .......................................................................................................................................................4

Cnidaria................................................................................................................................................................................................ 4

REPRODUCTION .............................................................................................................................................................4
Asexual .................................................................................................................................................................4
GENERAL ASPECTS .........................................................................................................................................................4
MORPHOLOGICAL FORMS ................................................................................................................................................5
Polyp .....................................................................................................................................................................5
Medusa .................................................................................................................................................................5
NERVOUS SYSTEM ..........................................................................................................................................................5
CUBOZOA .....................................................................................................................................................................5
ANTHOZOA ...................................................................................................................................................................6

Ctenophora......................................................................................................................................................................................... 6

GENERAL ASPECTS .........................................................................................................................................................6

REPRODUCTION .............................................................................................................................................................6

Phylogenetic Aspects of Phyla studied so far ............................................................................................................................ 6

Clade Lophotrochozoa .................................................................................................................................................................... 6

Phylum Platyhelminthes................................................................................................................................................................... 7

GENERAL ASPECTS .........................................................................................................................................................7

NERVOUS SYSTEM ..........................................................................................................................................................7
OSMOREGULATION & EXCRETION .....................................................................................................................................8
REPRODUCTION AND REGENERATION .................................................................................................................................8
CLASSIFICATION .............................................................................................................................................................8
Class Turbellaria ...................................................................................................................................................8
Class Trematoda ...................................................................................................................................................8
Class Monogenea .................................................................................................................................................8
Class Cestoda ........................................................................................................................................................8
EVOLUTIONARY ASPECTS .................................................................................................................................................8

Phylum Annelida ............................................................................................................................................................................... 9

GENERAL ASPECTS .........................................................................................................................................................9

BAUPLAN .....................................................................................................................................................................9
NERVOUS SYSTEM ..........................................................................................................................................................9
CLASSIFICATION .............................................................................................................................................................9
Polychaeta ............................................................................................................................................................9
Oligochaeta ..........................................................................................................................................................9
EVOLUTIONARY SIGNIFICANCE OF THE COELOMIC FLUID & METAMERISM ...............................................................................10
ADAPTIVE DIVERSIFICATION ...........................................................................................................................................10
EVOLUTIONARY ASPECTS ...............................................................................................................................................10
Phylum Rotifera ................................................................................................................................................................................10

BODY PLAN.................................................................................................................................................................10
REPRODUCTION ...........................................................................................................................................................10
EVOLUTIONARY ASPECTS ...............................................................................................................................................11

Clade Lophophorata ....................................................................................................................................................................... 11

Phylum Ectoprocta ........................................................................................................................................................................... 11

REPRODUCTION ...........................................................................................................................................................11

Phylum Mollusca ..............................................................................................................................................................................12

EVOLUTION OF THE TRUE COELOM ..................................................................................................................................12

GENERAL ASPECTS .......................................................................................................................................................12
BAUPLAN ...................................................................................................................................................................12
Mantle ................................................................................................................................................................12
Head-Foot ...........................................................................................................................................................12
Mantle Cavity .....................................................................................................................................................12
Shell ....................................................................................................................................................................13
Circulation and Gas Exchange ............................................................................................................................13
Other Visceral Modules ......................................................................................................................................13
Reproduction ......................................................................................................................................................13
CLASSIFICATION ...........................................................................................................................................................13
Class Monoplacophora .......................................................................................................................................13
Class Polyplacophora ..........................................................................................................................................13
Class Gastropoda ................................................................................................................................................14
Class Bivalvia (Pelecypoda/Acephala) ................................................................................................................14
Class Cephalopoda..............................................................................................................................................15
Porifera
Reproduction
All capable of sexual reproduction, w/ several types of asexual processes common.

Gametes & embryos occur throughout the mesohyl.

Asexual:
Gemmules: produced by freshwater Spongillidae in winter

Sexual:
Most sponges are hermaphroditic.

Sperms arise from choanocytes, eggs arise from choanocytes/ archaeocytes.

Diplontic life cycle

Sperm release looks like plant pollination

Most viviparous (w/ internal fertilization), choanocytes recognize and transfer (transcytosis) sperms of the same
species

Some oviparous (w/ external fertilization)

Larva types:
Parenchymula: embryo released lately, solid, w/ an outer surface of monoflagellated cells and inner mesohyl-like
core of matrix and cells [most demosponges]

Coeloblastula: embryo released early, two types of development, [calcarea, a few demosponges]:

1. inward migration of cells w/o flagella, they then regain it

2. stomoblastula> amphiblastula: stomoblastula has two cell types: micromeres and macromeres, constituting
separate poles, and is hollow. Inversion then occurs, moving the flagellated cells from the outside to the inside,
resulting in amphiblastula larva (macromeres> pinacocytes, micromere> choanocytes)

General aspects
Sponges in calm waters grow taller than those in rapidly moving waters. Generally, sponge development is highly
adaptable, depending on substrate and space.

The absence of organs means that basic processes such as excretion and gas exchange all occur at the cellular
level, mediated by diffusion.

Calcarea have all three body forms, hexactinellida have syconoid or leukonoid body form, and all demospongia
are of the leuconoid type.

Skeletal framework
The framework can be fibrous and/or rigid. When rigid, it consists of calcareous/siliceous spicules. The fibrous part
is present in all sponges and consists of collagen, one form of which is called spongin.

Most asconoid sponges are small because of the innate low surface-to-volume ratio of their spongocoel
In the syconoid sponges, food capture doesn’t occur in the spongocoel, so it’s lined w/ epithelial-type cells rather
than choanocytes.

In the leukonoid sponges, the small size of the flagellated chamber permits efficient filtering because of high
surface-to-volume ratio.

Cell types
Amoebocytes:
Move through the mesohyl, various functions

Take up nutrients from the surrounding water and choanocytes, digest it, and may carry it to other cells

Also secrete skeletal fibers in the mesohyl

Capable of diff. into other types of sponge cells

Cnidaria
Diverse aquatic habitats, some symbiotic.

Freshwater hydras and reef-building corals bear symbiotic algal cells in their tissues. This limits the occurrence of
reef-building corals to relatively shallow areas where there is sufficient oxygen available.

Reproduction
Asexual
By budding or fission

In budding, a knob of tissue forms on the side of an existing polyp and develops a functional mouth and tentacles.
If a bud stays attached to the polyp that made it, a colony will form and food may be shared through a common
gastrovascular cavity. So a budding polyp is expected to be colonial, and vice versa.

Fission common in Anthozoa.

General Aspects
Some w/ muscular contractions using epitheliomuscular cells, which form an outer layer of longitudinal fibers at
base of epidermis and an inner layer of circular fibers at base of gastrodermis.

Sense organs include well-developed statocysts (organs of balance) and ocelli (photosensitive organs); complex
eyes in members of Cubozoa.

Two nerve rings in hydrozoan medusae.

Sexual reproduction by gametes in all medusae and some polyps; monoecious or dioecious; planula larval form.

No excretory or respiratory system; No coelomic cavity.

(Important) A shared gastrovascular cavity permits specialization of the zooids, e.g. for reproduction, protection,
feeding, etc. For example in class hydrozoa hydranths are easily distinguished from the usually medusae-
producing gonangia by the latter’s lack of tentacles.
Cnidarian bodies extend, contract, bend, and pulse, all in the absence of true mesodermally derived muscle cells.
Epitheliomuscular cells form the functional equivalent of a layer of longitudinal muscle next to the mesoglea.
Contraction of these fibrils shortens the body or tentacles.

The mesoglea is gelatinous, or jellylike, and both epidermal and gastrodermal cells send processes into it. In
polyps, it is a continuous layer extending over both body and tentacles, thinnest in the tentacles and thickest in
the stalk portion.

The mesoglea is much thinner in the bells of hydromedusae and lacks amoeboid cells or fibers; water in the
gastrovascular cavity serves as a hydrostatic skeleton

In Hydra, the tentacles are hollow and the tentacle cavity communicates with the gastrovascular cavity.

Corals supplement their nutrition by collecting carbon from their algal symbionts.

The gastrovascular cavity is continuous from mouth to tentacles, and the gastrodermis lines the entire system
(hence the name gastroVASCULAR)

Cnidocytes occur throughout the epidermis. Hydras have three functional types of cnidae: those that penetrate
prey and inject poison (penetrants), those that recoil and entangle prey (volvents), and those that secrete an
adhesive substance used in locomotion and attachment (glutinants).

Morphological Forms
Polyp
The base by which colonial hydroids attach to the substratum is a rootlike stolon, or hydrorhiza.

Medusa
Free swimming, bell or umbrella-shaped bodies often w/ tetramerous symmetry.

Hydromedusae differ from scyphomedusae by the presence of a velum, a shelf-like fold of tissue from the bottom
of the bell that extends into the bell. By reducing the cross-sectional area at the bottom of the bell, the velum
increases the exit velocity of water from the bell, making each pulsation more efficient.

A life utilizing both a polyp and a medusa takes advantage of both benthic & pelagic environments, e.g. true jelly-
fishes and most hydroids of class Hydrozoa. However, many variations exist, such as polyp colonies drifting in the
ocean, as in Physalia, which use an inverted polyp as a gas-filled float (#Sari actually interpreted as colonial
Medusae)

Nervous System
In scyphomedusae and the medusae of cubozoans, nerves are grouped in marginal sense organs, called rhopalia,
that house chemoreceptors, statocysts, and often ocelli. The nerve nets form two or more systems, including a
fast-conducting system to coordinate swimming movements and a slower one to coordinate movements of
tentacles.

The nerve net arose early in metazoan evolution, and it has never been completely lost phylogenetically. Annelids
have it in their digestive systems. In the human digestive system it is represented by nerve plexuses in the intestine
musculature. Rhythmical peristaltic movements of the stomach and intestine are coordinated by this counterpart
of the cnidarian nerve net.

Cubozoa
Pedalium at the base of the tentacles, velarium which is similar to velum but different structurally.
Anthozoa
The mesoglea is a mesenchyme containing amoeboid cells.

A general tendency toward biradial symmetry in the septal arrangement occurs in the shape of the mouth and
pharynx. (Note how the tendency of stimuli to come from one direction (pharynx) causes bilateral symmetry)

Ctenophora
General Aspects
8 rows of comb-rows for locomotion, each w/ transverse comb-plates. Mostly free-swimming. Biradial symmetry
results from to tentacles on the body, w/ oral-aboral axis.

Tentacles used in pray capture, but in ctenophores w/ short tentacles, food is collected on the ciliated body
surface.

Colloblasts: adhesive structures on tentacles

Statocysts at the aboral end, surrounded by tufts of cilia

Most members emit flashes of luminescent at night

The middle gelatinous layer contains muscular fibers of various orientations

Reproduction
Monoecious w/ external fertilization

Free-swimming cydippid larva

Phylogenetic Aspects of Phyla studied so far

Both phyla clearly diploblastic, but the presence of cell in the mesoglea is problematic. If their origin is
endodermal, then they are somewhat homologous to a true mesoderm, if ectodermal, then they are derived
independently.

Neither phyla has a major deviation from the basic body plan.

Porifera are considered to be evolutionary dead-ends, giving rise to no other animal phyla.

Clade Lophotrochozoa
United by molecular similarities, share either of two morphological synapomorphies, an odd horse-shoe shaped
feeding structure called the lophophore (seen in Ectoprocta & Brachiopoda) or a particular larval form called the
trochophore. Trochophore larvae occur in many marine members of Annelida & Mollusca, as well as some marine
members of Platyhelminthes, Nemertea, Echiura, Sipunculida, etc.
Phylum Platyhelminthes
Contain free-living forms (most Turbellaria, w/ cellular ciliated epidermis) as well as parasitic forms (Neodermata
w/ syncytial body covering, called the tegument). Monogenea are mostly ectoparasitic, w/ a single host,
Cestoidean (tapeworms) and Trematoda (flukes) are all endoparasitic, w/ multiple hosts. The intermediate host is
usually an invertebrate (notably a mollusk), and the definite host is usually a vertebrate.

Campbell divides them into two groups, Rhabditophora and Catenulida.

Highly successful group, especially as parasites.

Acoelomate animals are a non-monophyletic group.

General Aspects
The region between the digestive cavity and the epidermis is filled w/ a solid, mesodermally derived parenchyma.
Its difference w/ the mesoglea of Radiata is that it has more cells & fibers, and less ECM.

Typical acoelomate animals have a gut cavity lined w/ endodermal cells and surrounded by mesodermal tissue.

Cellular epidermis contains rod-shaped Rhabdites, which swell and form a protective mucous sheath around the
body when discharged w/ water.

The benefit of the Neoderm is that it’s more resistant, because there are no penetrable junctions between cells.
This is particularly useful for gut parasites.

Parenchyma cells are mostly not distinct cell types, but non-contractile portions of muscular cells.

The gut is blind, usually w/ blind branches as well (e.g. 1 anterior and 2 posterior branch in tricladida, and Y-shaped
branchings in Trematoda and Monogeneans).

Shedding of the epidermal layers of the embryo to encompass the ectolecithal yolk in the Neodermata may be
the evolutionary origin of the neodermis.

Nervous System
Contains a pair of anterior ganglia and longitudinal nerve cords, connected by transverse nerves located in the
mesenchyme in most members.

The most primitive system in this phylum is the subepidermal nerve plexus found in some Turbellaria. Other
members have, in addition to the plexus, 1-5 pairs of longitudinal nerve cords. Transverse connecting nerves result
in a ladder-like system.

Active locomotion has also favored elaboration of sensory organs, incl. light-sensitive ocelli, and ear-like structures
on the sides of the head (auricles) as well as statocysts and chemoreceptors. Sensory organs more concentrated
around certain openings, such as the oral sucker of trematodes and the scolex of cestoda, and genital pores in
both groups.

The phylum representative ladder-like nervous system is actually exclusive to the tricladida. The nervous system
of polycladida is pretty much net-like (note the gastrovascular anatomy of the polycladida and its relation to the
nervous system arrangement)
Osmoregulation & Excretion
Osmoregulatory systems remove excess water from the body, excretory systems remove nitrogenous wastes
from the body. When waste products are dissolved in water, these two systems are said to be combined.
Flatworms have protonephridia, which removes only a small portion of nitrogenous wastes (ammonia), because
most wastes are excreted by diffusion through the body covering. Protonephridia are most elaborate in
freshwater species.

Reproduction and Regeneration

Both asexual and sexual. Asexual reproduction by fission.

Regarding sexual reproduction, most members are hermaphroditic, but practice cross-fertilization.

Classification
Class Turbellaria
Most have locomotive dual glands at the ventral surface of the body.

Class Trematoda
Undergo fission in mollusk host and sexual reproduction in the definite host.

Most members w/ oral and ventral suckers.

A great portion of the body is devoted to reproduction, w/ multiple reproductive glands.

Schistozoma (blood flukes) are unusual in being dioecious, w/ the males being heavier and having a gynocephoric
canal to embrace the slender female.

Class Monogenea
Share a posterior attachment organ w/ the Cestoda.

Class Cestoda
Have excretory and nervous systems. No special sensory organs, but do have sensory endings in the tegument.

Their entire surface is covered w/ microtriches, similar in structure and surface-increasing role to the vertebrate
microvilli.

The youngest proglottids are those proximal to the scolex.

Some self-fertilize.

The scolex is the evolutionary remnant of the posterior part of the ancestral body.

Evolutionary Aspects
First phyla in this document w/ organ-level organization.

The branching of the gastrovascular cavity causes more surface-to-volume ratio, decreasing diffusion distance.
(Note that the gastrovascular cavity is also a vascular cavity!)

The evolution of protonephridia allowed the colonization of land.

Phylum Annelida
General Aspects
Their most important apomorphy is metamerism, and its importance is demonstrated w/ the diversity of the three
metameristic phyla combined. Each repeated unit consists of independent circulatory, nervous(?), and excretory
systems. This independence provides a safety factor; to maintain the function of the other segments in case one
fails.

Bauplan
Another apomorphy is the annelid head, consisting of two parts, a prostomium and a peristomium.

w/ hydrostatic skeleton in each segment; the volume of the fluid in each segment is constant, contraction of the
muscles changes the proportions of the segment.

Non-chitinous cuticle

The gut is not segmented, probably because the independent feeding of the segments would be problematic.

Respiratory pigments in plasma, amoeboid cells in plasma

Clitellum: ring of secretory epidermal cells, appears as a fat band about one third of the animal’s length from the
anterior end. (Permanent in Oligochaeta, present during the reproductive season in Hirudinea). Both classes in
clade Clitellata are monoecious animals.

Nervous System
Ventral nerve cords w/ segmental ganglia and cerebral ganglia

Classification
Polychaeta
Well-differentiated head, paired appendages called parapodia, no clitellum. Some sedentary.

Some w/ open circulation and loss or reduction of septae.

No permanent sexual organs, sexes separate.

Protonephridia or a combination of meta- and protonephridia

Oligochaeta
Antero-posterior infolding of the gut known as the typhlosole, w/ functions somewhat similar to the vertebrate
liver

Double transport system; use both the blood and coelomic fluid as circulatory fluids. The five first segments of
the dorsal blood vessel function as a true heart.

Use metanephridia for excretion. Aquatic species excrete ammonia, terrestrial species excrete urea
Evolutionary Significance of the Coelomic Fluid & Metamerism
The coelomic fluid, by acting as a circulatory fluid, makes numerous protonephridia unnecessary. Gametes could
be stored in the spacious coelom for later, simultaneous release during copulation. This raises the need for an
advanced nervous system and endocrine system.

The coelom may have evolved in accord w/ different needs and evolutionary pressures in protostomes and
deutrostomes.

The selective advantage of metamerism for annelids lies in the efficiency of burrowing made possible by shape
change in individual segments of the hydrostatic skeleton

Adaptive Diversification
Have undergone extensive adaptive diversification, such as numerous feeding structures, many functions of the
parapodia, and numerous changes in the leech body plan for its predatory or bloodsucking lifestyles.

Evolutionary Aspects
The functional significance of metamerism for annelids: flexible support & efficient locomotion.

Some speculate their proximity to arthropoda (metamerism, ventral nerve cord, complete digestive tract)

Phylum Rotifera
Body Plan
Syncytial epidermis w/ subepidermal muscles

Complete gut

Mastax w/ a muscular portion equipped w/ hard jaws (trophi)

Pair of protonephridia

Reproduction
Sexes separate, but no males in class Bdelloida; they have amictic eggs

Some sort of haplodiploidy practiced by class Monogononta; females mostly produce amictic eggs, but they may
produce mictic eggs in a variety of conditions. The mictic eggs, if not fertilized, develop into haploid males. If
fertilized, develop into diploid, amictic females. Males often do not grow and are often sexually mature by the
time of hatching.
During copulation, sperm are injected directly into the female’s pseudocoelom.

Evolutionary Aspects
One of the first phyla w/ the ability of mechanical breakdown of food. The complete digestive tract allows
sequential digestion, absorption, and feces formation to proceed continually in an anteroposterior direction.

Many members of the aschelminths (Rotifera, Nematoda, Priapula, etc.) exhibit eutely; i.e. having a fixed number
of cells (or nuclei in the case of syncytial organs such as tegument). They also have protonephridia, a cuticle, and
adhesive glands.

Clade Lophophorata
The tentacles on the lophophore have two functions: feeding, and respiration, since they’re hollow and contain a
portion of the coelom

All coelomate

Phylum Ectoprocta
Colonial (the members called zooids); mostly sessile, some slime. Zooids form an exoskeleton in which they live,
called a zoecium. The zoecium also has an operculum. The zooids may be modified for various functions, such as
avicularia, gonozooids, ovicells (brood chamber, for the development of the embryo) and gastrozooids.

Some might be easily mistaken for hydrozoan colonies, but the presence of an anus is good microscopic criterion.

Entoprocts have their anus in their tentacular crown.

U-shaped gut connected to their sessile lifestyle.

Reproduction
Mostly hermaphroditic
Phylum Mollusca
Evolution of the True Coelom
The problem w/ a pseudocoelom is that the organs lay loose in the body cavity. Whereas a true coelom is lined by
mesoderm (completely), and the organs are suspended by mesodermal membranes called mesentries.
Mesentries provided structural support for a variety of evolutionary breakthroughs incl. a complex network of
blood vessels, and a muscular, diversified and specialized alimentary canal.

General Aspects
Coelomate lophotrochozoan protostomes, spiral mosaic cleavage, coelom by schizocoely. Trochophore larva is
the ancestral condition, but it is variously modified in various groups. Some of the most sluggish AND some of the
swiftest invertebrates are in this phylum.

Incl. classes Caudofoveata, Solenogastres, Monoplacophora, Scaphopoda, Polyplacophora, Bivalvia (Acephala),

Cephalopoda, and Gastropoda.

Bauplan
Their coelom is not used in locomotion, so it is functionally quite different from the coelom of annelida.

The visceral organs depend primarily on ciliary tracts for their function.

Mantle
Two folds of the skin, outgrowths of the dorsal body wall, form a protective mantle; the mantle encloses a space
between the mantle and the body wall called the mantle cavity. The mantle houses the gills or lungs, and in some
groups secretes a protective shell.

Head-Foot
All molluscs except bivalves have a unique feeding structure within the mouth called the radula.

Various modifications of the foot for various forms of locomotion or attachment, incl. the attachment disk of the
limpets, the laterally compressed hatchet foot of the bivalves, and the siphon for jet propulsion found in squids
and octopuses. In bivalves and gastropods the foot is extended to the out of the body hydraulically, by
engorgement w/ blood.

Mantle Cavity
Of enormous importance, and numerous roles. Houses the gills or lungs, and its exposed surface also acts as a
respiratory medium; a continuous flow of water (either by ciliary beating or muscular pumping or swimming)
keeps the cavity ventilated. Products from the digestive, excretory, and reproductive system are emptied into the
mantle cavity. The same current that ventilates the mantle cavity also flushes out wastes and carries out the
reproductive products.

Many molluscs may withdraw their head and/or foot into the mantle cavity for safety.

The gill, in its simplest form, consists of long, flattened axis extended from the mantle cavity. Use countercurrent
exchange for more efficiency in gas exchange.
Shell
Consists of three layers. The outer, organic layer is called the periostracum and is synthesized only at the margin
of the shell and then becomes worn away at the oldest part. Used for protecting the inner layers from boring by
other organisms. The second layer, called the prismatic layer, consists of densely packed prisms of CaCO3 laid
down in a prt. matrix. The next layer is called the nacreous layer, is secreted continuously throughout the life of
the animal and thus increases in thickness as the animal ages.

Subject to many modifications. Thick periostracum protects against the acidic environment of fresh-water, and is
thin in marine species.

First appears during the larval period and grows gradually from then on.

Circulation and Gas Exchange

The ancestral condition is open circulation, consisting of vessels AND sinuses. Due to its inefficiency in gas
exchange, it is only common in slow moving animals (insects are a notable exception because of their tracheal
system)

Other Visceral Modules

Utilize a pair of metanephridia for excretion and osmoregulation.

Highly complex & specialized digestive tract.

The nervous system consists of several pairs of ganglia w/ connecting nerve cords; considered to be simpler than
that of annelids and arthropods.

Reproduction
Mostly dioecious, some hermaphroditic. The ancestral life history is a trochophore larva w/ direct metamorphosis
into a juvenile form, as seen in chitons. However, in many groups, especially gastropods and bivalves the
trochophore stage is followed by a unique molluscan larval form called veliger. It has the beginnings of a foot, shell,
and mantle. Many species, incl. all cephalopods, have no free-swimming larval stages and a juvenile hatches
directly from the egg.

Classification
Class Monoplacophora
Serially repeated organs. (Also seen to a lesser extent in Polyplacophora)

Class Polyplacophora
Mostly live in the intertidal zone

Flattened dorsoventrally. Convex dorsal surface bearing 7-8 plates.

Don’t get very far from their “home”.

The mantle forms a girdle over the shell, and may cover it completely in some species [cryptochitons]. Its cavity is
extended along the sides of the foot. w/ the foot and mantle margin adhering tightly to the substrate, the lateral
mantle cavities become closed chambers w/ openings only at the anterior and posterior ends. Water flows
anteroposteriorly to ventilate the cavity.

Nephrostomes are in the pericardium lumen.

Nerve ring around the mouth

Class Gastropoda
Univalve shell w/ or w/o coiling. The apex contains the smallest and oldest whorl, starting from there, the whorls
become successively larger and spiral around the central axis (columella). May be clockwise or counterclockwise.
Many shells have an operculum which covers the shell aperture when the body is withdrawn into the shell.

After torsion, the anus and mantle cavity end up opening above the head.

Lateral organs flip their positions, forming a figure eight arrangement.

Fouling, the emptying of the anus over the gills, is a curious evolutionary paradox regarding torsion. Several ideas
have been proposed for the adaptive significance of torsion, such as the mantle cavity sensory structures facing
the anterior end of the animal. Another significance might be a withdrawing the head first, not last, into the shell
as a defensive behavior.

Coiling resulted in the conversion of the ancestral planospiral shell to a conispiral one; to solve the compactness
problem of a planospiral shell. Balancing of the bilaterally asymmetric shell resulted in the loss of right kidney, gill,
and atrium.

Many feeding habits, from filter feeding to active predation. Some bore a hole into their bivalve prey’s shell, and
some use their poisonous proboscis like a harpoon, which is a significant adaptation for a slow moving predator.

Opisthobranches and prosobranches use ctenidia (gills) for respiration, two in ancestral and one in derived, coiled
conditions. Pulmonates use a highly vascularized area of the mantle called lung, which opens to the outside by a
small opening to the out of the mantle called a pneumostome. Contraction of the mantle floor ventilates the
mantle cavity.

Most members w/ a single nephridium (kidney)

The nervous system incl. three pairs of ganglia connected by nerves.

Incl. both dioecious and monoecious (requiring copulation) members. Fertilization internal in most members.

Different subclasses of gastropods use different methods for avoiding fouling. Opisthobranchia, for example,
have undergone detorsion, and prosobranchia use long siphons to separate excurrent and incurrent waters, and
pulmonates have a left to right direction of water flow for the gills.

Some sea slugs use photosynthetic organelles of their prey to photosynthesize.

Pulmonates also incl. most freshwater snails. Land forms have two pairs of tentacles w/ one eye on each tentacle
in the posterior pair.

Class Bivalvia (Pelecypoda/Acephala)

No radula, no head, very little cephalization (a result of their non-locomotive, burrowing life-style)

The umbo is the oldest part of the shell, and growth occurs in concentric lines around it.

Pearl production occurs when many layers of nacre are secreted around an irritating object between the mantle
and the shell.

Jet-propulsion locomotive forms can regulate the direction of the water current, allowing them to move in
virtually any direction.

The nervous system consists of three pairs of widely separated ganglia connected by nerves.

Sexes separate, gametes excreted through the excurrent siphon.

Class Cephalopoda
Modified foot concentrated in the head region

Shelled cephalopods incl. nautilus, squids have internal shells (pen) and octopuses don’t have any.

Septate chambers of the nautilus shell connected by a siphuncle.

Branchial hearts evolved because the molluscan circulation plan places the entire systemic circulation before the
gills (unlike the vertebrate single circulation plan in fishes w/ places the gills immediately after the heart).

Since statocysts regulate eye motion, the eyes are always held in a horizontal position.

Quite capable of conditioning and observatory learning.

Some capable of color change, mediated by their chromatophores.

Sexes separate. No free-swimming (i.e. not in the egg) larva exists in cephalopods.

Octopoda don’t have any tentacles.