Sponges (Porifera, Poriferans)

General Features
Sponges are either asymmetrical or exhibit fractal (branching) or radial symmetry. They
are at the cellular grade of construction and have no organs (as such), no mouth or
digestive tract and no nervous tissue. Radial forms have vase-like shapes, but most
sponges are irregular or plant-like and of modular construction, forming flat, rounded or
branching structures.
The sponge body is permeated by pores, canals and chambers through which a water
current flows. Numerous channels permeate the interior; some or all of which are lined
by choanocytes (collar cells). The incurrent openings (ostia) are numerous and small,
whilst the excurrent openings (oscula) are few and large. The choanocytes resemble
choanoflagellates and possess a single flagellum encircled at its base by a protoplasmic
collar.
Most sponges have an internal skeleton, secreted by mesenchymal amoebocytes and
composed of separate crystalline bodies (spicules) or organic fibres, or both, and may
incorporate foreign particles. The spicules are made of carbonate in some sponges and of
silicate in others.
All sponges are sessile and the adults are incapable of locomotion, and live fastened at
their base to rocks, shells, timbers, corals, plants, etc. Some sponges are not attached and
roll about freely on the bottom. Sponges are marine, except for the family Spongillidae
that lives in ponds and lakes throughout the world. As sponges grow they spread or
branch in a plant-like manner.
External Features
The Calcarea and the Hexactinellida contain the smallest and simplest sponges and have
radial symmetry. The body is cylindroid, vase-like and hollow and fastened at one end,
whilst narrowing to a single osculum at the other end. Many Choristida and Hadromerina
(Demospongiae) are more-or-less spherical with an internal radiating structure. Most
sponges, however, are irregular or have fractal symmetry. Sponges vary in size from
small to large, and are sometimes cup-shaped, funnel-shaped, or fan-like (flabelliform),
but are more often massive, encrusting and irregular or branching structures.
The sponge surface is studded with numerous pores and many larger oscula. The oscula
are often borne on the ends of branches or projections or sunk into crater-like
depressions. The form of sponges is highly dependent on the substratum and
environmental conditions (especially water currents).
All sponges are sessile and the adults are incapable of locomotion, and live fastened at
their base to rocks, shells, timbers, corals, plants, etc. Some sponges are not attached and
roll about freely on the bottom. Sponges that live on muddy bottoms in deep waters, e.g.

the hexactinellids, are fastened to the mud by root bundles of long spicules. Sponges may
be stalked.
Sponges vary in height/diameter from a few mm to large rounded masses or upright
growths of 1-2 m. Calcareous and siliceous sponges have a bristly and rough exterior, due
to the presence of projecting spicules. Horny sponges and some other sponges, however,
are slimy or smooth, or have a hard, leathery surface.
Most sponges are of a drab, flesh, or brownish colour, but some are bright orange, yellow,
red, blue, violet or black, due to the presence of lipochrome pigment in some of the
amoebocytes. Deep-water forms tend to be drab, whilst shallow water forms tend to be
more brightly coloured.
Fresh-water sponges are brownish-green due to zoochlorellae. These sponges are
encysting or slightly branching and attached to submerged objects and have monaxon
siliceous spicules.
Structure
1. The asconoid is the simplest structural type of sponge. The asconoid has radial
symmetry and is vase-like. A thin wall surrounds the large central cavity, or spongocoel,
which opens at the summit via a single narrowed osculum. The wall consists of an outer
monolayer of epidermis of thin, flat cells, and an inner epithelium comprised of a singlelayer of choanocytes in loose contact with each other. A layer of mesenchyme
(ectomesoderm) is sandwiched between these two epidermal layers and consists of
amoebocytes and spicules embedded in a gelatinous matrix.
Numerous incurrent pores, the ostia, extend from the external surface to the spongocoel
as intracellular pores; each forming a canal through a tubular cell called a porocyte. The
choanocyte flagella generate a water current drawing water through the ostia and into the
spongocoel and out through the osculum. This water current supplies the sponge with
food and oxygen and removes waste.
The asconoid wall is simple with a complete and continuous choanocyte lining,
interrupted only by porocytes. Very few sponges exhibit the asconoid structure.
2. The syconoid is a more complex structure formed by outpushings of the asconoid wall
at regular intervals as finger-like projections or radial canals. There are two syconoid
structural subtypes. In subtype I these projections may be free and surrounded on the
outside by water along their whole length, in which case there are no definite incurrent
channels. Alternatively, in subtype II the projections may fuse to form incurrent canals,
tubular spaces between adjacent radial canals and the exterior has an epidermal lining.
These incurrent canals open to the exterior between the blind outer ends of the radial
canals via dermal pores or dermal ostia. The radial canals have a choanocyte lining and
form, flagellated canals. The large spongocoel is lined by flat epithelium derived from the
epidermis. The internal ostia are the openings of the radial canals into the spongocoel.

Syconoids are radial and vase-like with a single terminal ostium. The prosopyles are
pores between the incurrent and radial canals and may correspond to the ostia of
asconoids (intracellular? Apparently the prosopyles are intercellular spaces).
Flow of water is in through the dermal pores incurrent canals prosopyles radial
canals internal ostia spongocoel osculum outside.
Syconoid subtype I includes a few heterocoelous calcareous sponges, e.g. Sycon. In
subtype II the epidermis and mesenchyme are spread over the outer surface to form a
cortex (thin or thick) which may contain special cortical spicules and has definite dermal
pores. The dermal pores open into either narrowed incurrent canals that have an irregular
course and may branch and anastomose or into subdermal spaces that are the outer ends
of the flagellated canals.
3. The most complex structural type of sponge is the leuconoid type. This is formed by
continued outfolding of the choanocyte layer. The choanocytes are limited to clusters of
small, round or oval chambers. Mesenchyme fills the spaces around the flagellated
chambers. There is usually no spongocoel. Leuconoids have an irregular, indefinite form
permeated by a maze of water channels.
The surface of the leuconoid sponge is usually covered by epidermis (not in
hexactinellids) that is pierced by dermal pores and oscula. The flow of water is in through
the dermal pores into either incurrent passages (branching irregularly through the
mesenchyme) or into large subdermal spaces crossed by columns of spicules. Both
incurrent passages and subdermal spaces open via prosopyle openings into small,
rounded flagellated chambers. Water exits the flagellated chambers via the apopyle
apertures into the excurrent channels that unite into larger and larger tubes that
discharge to the outside through the oscula.
There are three structural subtypes. In the eurypylous system the apopyles open directly
via wide mouths into the excurrent channels. In the aphodal system a narrow canal, or
aphodus, connects the flagellated chamber to the excurrent channel. In the diplodal
system, a narrow tube (prosodus) connects the incurrent channel and the flagellated
chamber and there is also an aphodus connecting the chamber to the excurrent channel.
Often, the leuconoid body has an outer ectosome region devoid of flagellated chambers
and an inner choanosome (endosome) containing clusters of flagellated chambers. The
ectosome comprises a cortex or dermal membrane and subdermal spaces. The cortex
differs from the choanosome in histology and spicule type or spicule arrangement. The
dermal membrane consists of an epidermis and a thin mesenchyme stratum.
The leuconoid is characterised by the limitation of choanocytes to small chambers and
the great development of its mesenchyme and the complexity of its incurrent and
excurrent water passages.

Water flows in via the dermal ostia subdermal spaces and incurrent channels
prosodus if present prosopyle pores flagellated chambers apopyles aphodus
if present excurrent canals larger channels out via oscula.
The leuconoid sponge may develop through the asconoid and syconoid stages, but most
leuconoids develop from a rhagon stage. The rhagon is conical, tapering from a broad
base to a single osculum at the summit. The rhagon possesses a spongocoel bordered by
oval flagellated chambers opening into it by wide apopyles.
The majority of sponges are of the leuconoid type since this structure allows them to
generate a more efficient water current and to attain a larger size. The leuconoid form
probably evolved several times.
Histology
Sponges contain the following principal cell types: epidermal cells, muscle cells, gland
cells, choanocytes and mesenchymal amoebocytes.
Some sponges possess an epidermis of large, flat, polygonal epithelial cells called
pinacocytes. (Endo)pinacocytes line the incurrent canals and the spongocoel of
syconoids and the larger canals and spaces of leuconoids. Pinacocytes are highly
contractile and can greatly reduce the surface area of the sponge. Many sponges have a
syncytial epidermis (epithelioid membrane). However, the Hexactinellida have no
definite epidermis.
In asconoids, pore cells or porocytes are tubular cells pierced by an intracellular
incurrent canal or pore. Pinacocytes are highly contractile and can close the pore by
advancing a thin sheet of cytoplasm (pore diaphragm) from the edge to the centre at the
outer end of the canal. The dermal pores of syconoids and leuconoids are of a disputed
nature, but are closable. Freshwater sponges have dermal pores that begin as porocytes,
each of which disappears later to leave a pore in the epidermis. The prosopyles are
analogous to the incurrent pores of ascon sponges (but are they intracellular or
extracellular?)
The mesenchyme is a transparent gelatinous matrix (mesogloea) containing free
amoebocytes. The mesenchyme may be a collenchyma with few cells, or a parenchyma
with a high cell density. The amoebocytes are free to wander about the sponge and fall
into two main classes, lobopodous amoebocytes and colllencytes. Lobopodous
amoebocytes include pigmented chromocytes, thesocytes that store food reserves and
scleroblasts that secrete the skeleton. Scleroblasts are further divided into calcoblasts,
silicoblasts and spongioblasts, depending on the nature of the skeletal material secreted.
Collencytes have slender branching pseudopods and may form a syncytial network.
Archaeocytes are lobopodous amoebocytes each with a large nucleus, conspicuous
nucleolus and sometimes containing cytoplasmic inclusions. Archaeocytes are possibly

undifferentiated cells and produce the sex cells (as may choanocytes in some sponges?)
and are involved in regeneration since they can give rise to all other sponge cell types.
Gland cells put out long strands to the sponge surface and possibly secret slime
(amoebocytes also secrete slime).
Desmacytes (fibre cells) are found in layers in the cortex and around larger internal
channels, and are common in the Desmospongiae. Myocytes are fusiform muscle cells
that resemble smooth muscle cells and usually form a sphincter at the osculum and other
openings.
Choanocytes (collar cells) are rounded or oval cells with their base resting on the
mesenchyme. Their free end bears a transparent contractile collar encircling the base of
the long single flagellum. Choanocytes are larger in the Calcarea.
Skeleton
The mesenchyme secretes and contains the skeleton. The skeleton consists of spicules
and/or spongin fibres. The spicules (sclerites) are crystalline bodies and each is a spine
or a number of spines radiating from a point. Each spicule consists of an organic axis
surrounded by calcium carbonate or hydrated silica. Megascleres are the larger spicules
that from the main supporting framework. Microscleres are smaller flesh spicules strewn
throughout the mesenchyme. However, such a size distinction does not hold for
calcareous sponges and some other groups.
Spicules are classified according to the number of spines or axes as follows.
1. Monaxon spicules have a single axis, straight or curved. The style (monactinal
monaxon) results from growth in one direction only and is often rounded at one end
(strongylote end) and pointed (oxeate) at the other. Tylostyles are styles that have the
broad end knobbed. Acanthostyles are styles that are covered with thorny processes.
The pointed end of each style usually projects to the exterior of the sponge. Diactinal
monaxons (diactines, rhabds) result from growth in both directions from a central
point. Oxeas are pointed at each end, tornotes are lance-headed at each end,
strongyles are rounded at each end and tylotes have a pinhead like knob at each end.
Microscleres have the prefix micro-, e.g. microrhabds, microxeas, microstrongyles.
The Sigmatophora and Poecilosclerina have curved types of diactinal miscroscleres,
including C-shaped sigmas and bow-shaped toxas and chelas with recurved hooks,
plates or flukes at each end. Isochelas are chelas with both ends alike, whilst
anisochelas have unlike ends. Sigmaspires are spirally twisted sigmas. Streptasters
are short, spiny, microscleric monaxons and include spirally twisted spirasters, rodshaped sanidasters, plesioasters that have a few spines radiating from a very short
axis, and amphiasters that have spines at each end.
2. Tetraxons (tetractines, quadriradiates) have four rays radiating from a common
point. These rays are not in the same planes. Tetraxons may lose 1-3 of their rays.

Calthrops tetraxons have equal rays. Microcalthrops are microscleric calthrops.

Triaenes have one elongated ray, the rhabdome, and three short rays (the cladi or
clads) constitute the cladome. Developmental loss of one cladi results in a diaene
spicule. The cladome may be a simple or scalloped disc instead of separate cladi. If
such a disc occurs at both ends of the rhabdome, then the resulting spicule is a
birotular spicule or amphidisc. The triradiate or triactinal spicule is the most
common spicule type in calcareous sponges and is a variety of triaene in which the
rhabdome is lost to leave a cladome of three rays that lie almost in one plane.
3. Triaxon (hexactinal spicules) have three axes crossing at right-angles to give six
rays, some of which may be lost or reduced or branched or curved and may have
spines or knobs, etc. These spicules occur only in the class Hexactinellida.
4. Polyaxons have several equal rays that radiate from a central point. Asters are
microscleres (euasters as opposed to streptasters) and include oxyasters that have a
small centre and long pointed rays; strongylasters that have a small centre and long
rays with rounded ends; and tylasters that have a small centre and long rays with
knobbed ends. Polyaxons with a large centre and short rays are the spherasters, with
definite rays, and the sterrasters in which the rays are small projections from the
spherical surface.
5. Spheres form from concentric growth around the centre.
6. The desma is a megasclere formed from a minute monaxon, triradiate or tetraxon
spicule, forming the central crepis, upon which layers of silica are deposited. These
deposits develop branches and tubercles. Desmas are named after the crepis shape, as
monocrepid, tricrepid or tetracrepid. Desmas are usually united into a network to
form a reticulated skeleton (lithistid).
Spongin is a protein that forms a branching network. In the Monoaxonida, spongin often
binds the siliceous spicules together. In the Keratosa the skeleton consists entirely of
spongin (and embedded foreign particles).
Spicules are secreted by scleroblasts. This secretion process is best known for calcareous
spicules. A monaxon spicule begins inside a binucleate cell as an organic axial thread
between the nuclei, which is coated in calcium carbonate within a clear space inside the
scleroblast. The nuclei draw apart as the spicule lengthens and the cell separates into two
daughters. One daughter is the thickener and is situated at the outer end of the spicule,
which is usually the projecting end. The other daughter is the founder and is situated at
the inner end of the spicule and deposits most of the spicule as it moves inward, its
secretion is supplemented by that of the thickener. Upon completion of the spicule, both
cells wander off into the mesogloea (the founder leaves first).

Triradiate calcareous spicules are formed by three scleroblasts that come together in a
trefoil. Each scleroblast divides into a founder at the inner end and a thickener at the
outer end and three spicules are secreted joined together as the founders move towards
the tips. The three thickeners remain awhile at the junction as they thicken the spicule.
Quadriradiate spicules are formed by an additional scleroblast adding a 4th ray to a
triradiate spicule.
In contrast, siliceous spicules are probably formed entirely within one silicoblast or by
several silicoblasts in the case of larger spicules. Hexactinal spicules, found in the
Hexactinellida, arise in the centre of a multinucleate syncytial mass (formed from a single
silicoblast?) as silica is laid down in concentric cylinders around a conspicuous organic
axial fibre (called the protorhabd).
Spongin is secreted by spongioblasts that arrange themselves into rows and the spongin
rod segment secreted by one cell fuses with those of adjacent spongioblasts to produce a
long fibre. After secreting a certain amount of spongin, each spongioblast vacuolates and
degenerates.
Physiology Movement:
Adult sponges are immotile and exhibit limited movement. The only movements
exhibited by some sponges are changes in the porocytes. Most sponges, however, are
capable of local or general contraction due to forces produced by the pinacocytes,
desmacytes or the myocytes. Such contractions are especially noticeable in some ascons.
Most calcareous sponges exhibit slight or no contractility. Many Desmospongiae can
contract their whole surface as a result of contractions of fibre cells in the cortex and
along the main channels. Sphincters control the aperture sizes of the oscula. Dermal pores
can open and close as a result of sphincter action or changes in the encircling cytoplasm.
Physiology Sensitivity:
Sponges have no nervous system and no specialised sensory cells. Disturbances result in
general body contraction. Oscula close on exposure to air, in response to injury, lack of
oxygen, chemical irritation, extremes of temperature and sometimes also to touch. The
oscula also close when the sponge is in a small volume of water that is inadequate for
waste removal. The dermal pores are less responsive, but close on injury to the sponge.
Apparently neither of the behaviours of oscula, dermal pores and choanocytes is
correlated (?). Reactions are slow, taking place over one to several minutes. Obnoxious
stimuli are transmitted either not at all or at up to 3-4 mm maximum.
The osculum is the most conductive area, especially in the direction away from the
osculum, and the oscular rim is the most sensitive part. In the freshwater sponge
Ephydatia each osculum is borne on the summit of a chimney-like tube. If the oscular rim
is stimulated, a signal is transmitted down the chimney causing the chimney to contract
or collapse.

Physiology Canal System:

In Leucandra, the speed of the oscula excurrent has been measured at 8.5 cms -1, and
travels as a jet for up to 25-50 cm. A Leucandra sponge 10 cm high and 1 cm in diameter
has some 2.25 x 106 flagellated chambers and filters 22.5 l of water per day. On this basis
a large sponge would filter about 100 l per day. The flow rate inside the flagellated
chambers is very slow at about 0.01 mms-1 due to a slight pressure gradient regulated by
the sizes of the oscula apertures.
The flagella do not beat in coordination. Each undergoes a spiral undulation that travels
from the base to the tip and ensures that the water current travels in a single direction.
The collars point towards the apopyle and the resulting current exits through the larger
aperture. Smaller flagellated chambers are more efficient since they permit less
stagnation.
In leuconoids the water current is very slow in the flagellated chambers, allowing time
for filtering and exchange of materials to occur. The current then speeds up as it leaves
the sponge.
Physiology Nutrition:
Sponges are filter feeders and sieve out microorganisms and organic debris from the
water flowing through the flagellated chambers. Dissolved nutrients are possibly also
taken up directly by the sponge (?).
Food particles in the water current adsorb to or are filtered by the choanocyte collars and
ingested into the cytoplasm. In the large choanocytes the particles maybe wholly
digested, but are usually either partially digested or passed straight onto the amoebocytes
of the mesenchyme for intracelluler digestion. The smaller choanocytes of other sponges
possibly have no digestive role but pass food directly onto the amoebocytes (?). It is also
possible that amoebocytes receive some particles absorbed directly through the walls of
the incurrent passages (?). The amoebocytes eject waste that leaves the sponge via the
excurrents.
Physiology Respiration:
Sponges are aerobic. Sycon consumes 0.16 0.04 cm3O2/g/fresh-weight/h, larger
specimens consuming less oxygen per unit mass than smaller ones, as is the case for
animals generally. Oxygen consumption is reduced by 80% when the oscula are closed
and this is compensated for by supernormal oxygen consumption when the oscula reopen.
Physiology Excretion:
Ammonia is excreted and waste-laden amoebocytes are possibly discharged (?). Some
sponges exude mucus or slime that has an unpleasant odour and may cause irritation to
the skin on contact.

Ecology
Many animals live on or in sponges, exploiting their sessile habit. Coelenterates,
bryozoans and barnacles may grow on the surface of sponges, whilst annelids and
crustaceans often inhabit the water canals. In one large specimen of loggerhead sponge
(Spheciospongia) were found 16 352 Synalpheus shrimps. Shrimps of the family
Stenopidae, especially Spongicola, live in pairs in the spongocoel of hexactinellid
sponges, such as Euplectella and Hyalonema. The shrimps enter the sponge when young
and grow such that they can no longer escape through the oscular sieve plate.
Sponges of the family Suberitidae grow on the snail shells inhabited by hermit crabs. The
sponge grows to enclose the shell, the shell is dissolved and the crab occupies a spiral
cavity in the sponge, which is lined by smooth fibrous tissue. The larvae of this sponge
possibly only grow when they land upon a shell containing a hermit crab (?).
Freshwater sponges (and marine?) may be parasitised by mites. The female mites lay
eggs in the sponges tissue.
Crabs, such as Dromia and others, break off pieces of living sponge (and other objects)
and hold them over their backs with their last pair of legs, for camouflage. Some crabs,
e.g. F. Majidae, stick sponges, algae and hydroids onto their back and legs with an
adhesive secretion. These implanted organisms may continue to grow.
Sponges are seldom eaten since they are prickly and have a bad taste/odour. Some
crustacean inhabitants feed on sponge tissue as parasites. Some nudibranchs also eat
sponges.
Sponges may smother and kill other sessile animals, as they grow, including oysters.
Boring sponges may kill barnacles and other shelled animals.
Large horny sponges live for 50+ years. Smaller sponges live for several months or years.
Freshwater sponges are necessarily seasonal.
Different sponge morphologies suite different waters. Sponges in calm or silty water have
elevated oscula. Sponges that live in rough waters are low and encrusting. Flabellate /
lamellate sponges thrive in water with a constant current, and grow with their inhalent
surface facing the current and their oscula point downstream. Deep-water sponges living
on muddy bottoms have anchoring root tufts or long projections of spicules. Most marine
sponges live in shallow water, from the tidal zone down to 50-m depth, and do not live in
brackish water. The hexactinellids are deep-water sponges. Some Demospongiae live in
the deep ocean oozes. Spongin skeletons prevail in tropical / subtropical waters, whilst
mineral skeletons are more abundant in cold waters.

Reproduction
Asexual Reproduction. Some sponges constrict off the ends of branches, which then
round up into a ball and regenerate. This may occur regularly or under adverse
conditions.
All freshwater, and some marine, sponges form gemmules. A gemmule is a group of
amoebocytes (archaeocytes?) with other amoebocytes forming a covering columnar layer
that secretes thick and hard inner and outer membranes. The amoebocytes receive glycoor lipoprotein food reserves from special nurse cells (trophocytes). Scleroblasts deposit
radial amphidisc spicules between the membranes (not in Spongilla). The result is a
round ball of archaeocytes, trophocytes and columnar cells, with a micropyle outlet.
The freshwater sponges produce gemmules in autumn and then die. The gemmules are
able to resist winter drying and freezing and hatch in spring. Upon hatching, cells stream
out through the micropyle and form a young sponge in about one week.
In marine gemmule producing sponges, the gemmule consists of an amoebocyte
aggregation (archaeocytes?) with a flagellated columnar dermal covering and the
gemmule becomes a free-living flagellated larva. The larva attaches near its posterior
nonflagellated pole, loses its flagella, and develops into a young sponge.
Sexual Reproduction. All sponges can reproduce sexually. Ova and spermatozoa are
produced, possibly from archaeocytes, other amoebocytes or from choanocytes (?). The
egg mother cell (ovocyte) is an enlarged amoebocyte that grows either by engulfing other
amoebocytes or from nutrients supplied to it by trophocytes. And then undergoes
maturation divisions.
Each sperm mother cell is an enlarged amoebocyte, or possibly a transformed choanocyte
since whole flagellated chambers have been observed to transform into spermatozoa. The
sperm mother cell becomes covered by one or more flattened covering cells, which derive
from divisions of the sperm mother cell or from other amoebocytes. The whole structure
is called a spermatocyst. The enclosed spermatogonium undergoes 2-3 divisions to
produce spermatocytes.
The sperm enter the recipient sponge in the water current and fertilisation is internal. In
the Calcarea, the sperm enters a choanocyte nurse cell. The nurse cell then fuses with the
egg, releasing the contained sperm. In Reniera the sperm enters an amoebocyte, which
transfers the sperm to the egg.
The fertilised egg undergoes unequal holoblastic cleavage to produce a blastula, in situ.
The blastula is flagellated and exits the sponge via the osculum. This larva swims for
several hours before attaching and developing into a sponge.

Most sponges are hermaphroditic, but some are dioecious. Hermaphrodite individuals
often produce eggs and sperm at different times, whilst some produce sperm apically and
eggs basally. Shallow water forms reproduce seasonally.
Regeneration
Sponges have very high regenerative powers. Any piece can regenerate into a whole
sponge, but the process is slow, requiring months or years to reach full size. If a sponge
broken into cells and cell-clumps, then the amoebocytes aggregate to form a reunition
mass. Some reunition masses contain collar cells without collars and various types of
amoebocyte. Some of the amoebocytes form epidermis and a whole sponge is reformed.
Reunition masses composed entirely of choanocytes cannot reform a sponge. Cells from
different species may temporarily form a reunition mass before separating.
In adverse conditions, many marine sponges and freshwater sponges, collapse and
disintegrate to leave a reduction body remnant comprised of a covering epidermis and an
internal amoebocyte mass with partially dedifferentiated choanocytes. This will grow into
a sponge when and if favourable conditions return.
Class Calcarea (Calcispongiae)
Calcareous sponges are small, up to 15 cm in height, and mostly drab in colour and often
have a bristly texture due to the presence of projecting monaxons. These sponges are
highly individualised and are solitary or else live in groups of radially symmetric vaselike bodies, each with its own terminal osculum. Some form bushy branching colonies, or
compact masses. Others are borne on slender stalks. The sponge is fastened to a hard
substrate at its base.
The osculum is often encircled by a collar of upstanding long monaxon spicules that form
an oscular fringe. A sieve membrane may cover the osculum. The spicules are made of
calcium carbonate and are usually separate, but are fused into a network in pharetrone
sponges, or are enclosed in calcareous cement. All the spicules are megascleres and
include monaxons (styles and rhabds) and triradiates, quadriradiate spicules (often oxeas)
are sometimes also present. However, triradiate is the dominant spicule type. T-shaped
sagittal triradiates interlace to form the oscular rim. If a cortex is present, then it contains
spicules parallel to the surface that form a strong layer.
Calcarea includes all asconoid forms and also some syconoid and leuconoid sponges.
Most of the ascons belong to the genus Leucosolenia. All the ascons form colonies of
tubes, united by horizontal tubes or a network of branching tubes or else the outermost
tubes fuse to give an apparently solid mass with a false surface (pseudoderm) permeated
by a few large pseudopores.
Clacarea also includes all typical syconoids, e.g. Sycon, Sycetta and Grantia. These
usually form single vases or clusters of vases, each with a large central spongocoel and a

terminal osculum that may have an oscular fringe. Syconoid forms with a cortex include
Grantiopsis, Grantessa, Heteropia, Sycaltis, Ute and Amphoriscus.
Calcareous leuconoids are all eurypylous and include Leuconia, derived from the
syconoid form, and Leucetta, which is derived from the asconoid form. The class also
includes syconoid-leuconoid transitional stages such as the sylleibid type sponges:
Rhabdodermella, Leucilla and Vosmaeropsis. There are also some asconoid-leuconoid
transitional forms resulting from anastomosis of ascon tubes and formation of
pseudoderm, as in Leucaltis (Heteropegma) and Leucascus.
In Sycon, the skeleton consists of layers of spicules (a so-called articulate skeleton). The
family Amphoriscidae (inc. Amphoriscus, Leucilla, Rhabdodermella) the skeleton is
inarticulate and consists of large triaenes or sagittal triradiates. Triradiate spicules have
three rays, with equal angles in regular triradiates. In sagittal or alate triradiates, the
paired lateral rays are at an angle < 120o and the third ray is the posterior or basal ray,
resulting in T-shaped or Y-shaped spicules. Some with their basal ray directed towards the
spongocoel (more or less spanning the cortex) and their laterals parallel to the surface,
others with the opposite polarity (basal rays directed towards the cortex). Triradiates form
the bulk of the skeleton in most Calcarea and are strewn throughout the interior in
leucons.
There is no gemmule formation in the Calcarea. Reduction bodies may form in adverse
conditions. Asexual reproduction by budding is common and breaking-off of branch tips
is a regular asexual reproductive mode in some ascons.
In sexual reproduction, the ova originate from archaeocytes. The embryo develops into a
stomoblastula that has a mouth with which it ingests adjacent maternal cells. By a process
of inversion (turning inside out as in the protoctistan Volvox) an amphiblastula larva
results. (During inversion the endoderm becomes the epidermis and the ectoderm
becomes the internal cells). The amphiblastula escapes to the outside via the water canal
system. The amphiblastula then either develops into a typical flagellated gastrula or into a
solid flagellated stereogastrula (parenchymula) larva. The (stereo)gastrula larva
attaches and develops into an asconoid olynthus stage and then into a syconoid type
sponge (and possibly a leuconoid form in some, either directly from the olynthus or via a
syconoid intermediate?). Some Leucoslenia develop via a free-living stereogastrula that
after a few hours attaches and flattens and develops into an asconoid sponge.
Class Hexactinellida (Hyalospongiae, Glass Sponges)
The hexactinellid skeleton consists of hexactine siliceous spicules, both megascleres and
microscleres. Most hexactinellids are strongly individualised, radially symmetric and
cylindrical, vase-, urn-, or funnel-shaped, etc. They are fastened at their base either
directly or via a root tuft of spicules. The spongocoel opens via an osculum at the summit
and there may be a sieve plate of silica covering the osculum or the osculum may be
bounded by upright spicules. Some hexactinellids are branching and some are lamellate.

Glass sponges are of a pale colour and projecting spicules gives them their characteristic
glass wool appearance. Most are 10-30 cm, but some are over 1m in length. All are
deep-water marine dwellers.
The internal structure consists of a trabecular net, which is a thin strand network
outlining large open meshes. This network is formed from amoebocyte pseudopods and is
apparently syncytial. There is no definite epidermis and the external surface, as well as
the spongocoel lining, are simply composed of the trabecular net and so is very open
with large openings. Hexactinellids are syconoid or simple leuconoid. The body wall
typically consists of the following regions: dermal surface, subdermal trabecular net, a
layer of flagellated chambers, a subgastral trabecular net and the spongocoel.
The hexactinellid skeleton is glass-like and some of the siliceous spicules are usually
loosely bound or fused into a lattice, while the rest are individual. The spicules consist of
silicate with a spiculin organic axis and sheath. The spicules are hexactines. The regular
hexactine has three axes at right angles to each other, giving six rays of about equal
length. Some have one elongated sword-like ray, and when this ray is covered in spines,
the spicule is called a pinule. Loss of one ray results in a pentact and loss of one axis
results in a tetract (cross or stauractine) with the 4 rays in one plane. Loss of three rays
results in the uncommon triact and loss of 4 rays results in the abundant diactine.
Uncinate spicules are diactines covered with short spines directed to one end. Loss of 5
rays results in the monactine. Clavules are monactines with a disc or bulb at one end,
scopules are monactines with a branched end and sceptres are monactines with little
spines along one end. The same types occur as microscleres and also as asters and
amphidiscs. The asters are regular hexactines with branched rays, the branches lacking
axial fibres. In oxyhexasters the ray ends divide into a few simple straight branches. In
discohexasters the branches are topped by discs and in discoctasters the branches form
brush-like bundles and are called plumicome if the brushes curve outwards and
floricomes if they have expanded tips. The amphidiscs are diactines ending in umbrellalike expansions.
The spicules of a hexactinellid sponge can be classified into several functional types.
Prostals are spicules that project from the sponge as long needles, and are mostly
diactines. Marginal prostals encircle the osculum and protrude singly or in bundles
(pleural prostals). Root tufts are called basal prostals or anchoring spicules. There are
recurved hooks on the distal ends of the root spicules for anchorage. Parenchymalia are
spicules strewn throughout the trabecular net. Dermalia are spicules at or just beneath the
dermal surface and gastralia are spicules at or just beneath the inner surface.
In early stages the network lattice is of the lyssacine type, in which the rays of hexactines
may be joined together by silica crossbars. The fully developed trabecular net is the
dictyonine type skeleton in which the rays are fused at their tips by layers of silica.
Hexactinellids are deep-sea sponges (100 m to 5000 m) and are mostly known from
preserved specimens. A gelatinous matrix appears to be absent. Myocytes surround some

of the openings, but otherwise these sponges are non-contractile due to their rigid
skeletons. The choanocytes appear to be fused basally into a syncytium.
Some hexactinellids reproduce asexually by budding, others by gemmule formation.
There are two orders of glass sponge, the Amphidiscophora, which have a lyssacine
skeleton and amphidisc microscleres and the Hexasterophora that have astrose
microscleres and usually have a dictyonal or a mixed lyssacine-dictyonine skeleton.
Example amphidiscophorans are Pheronema, Monoraphis and Hyalonema. Monoraphis
has a single gigantic anchoring spicule, which is 2-3 m long and 1 cm thick. In
Hyalonema the root tuft is often spirally twisted and continues through the sponge as an
axis or columella, and often projecting above the sponge as a gastral cone.
Examples of the Hexasterophora are Farrea, which has a quadrangular skeletal mesh,
Eurete with its triangular mesh and Aphrocallistes with a hexagonal mesh. Other
examples are Aulocystis, Euplectella, Caulophacus, Rossella, Rhabdocalyptus and
Staurocalyptus. Euplectella has a beautiful glassy skeletons form the Venuss flower
baskets, and possess a sieve plate, root spicules and a square lyssacine network.
Class Demospongiae
Demosponges either have no spicules at all or possess non-hexactinellid siliceous
spicules. The horny sponges have skeletons of spongin fibres. All are leuconoid (maybe
eurypylous, aphodal or diplodal) and derived from a rhagon stage. There are three
demosponge subclasses: Tetractinellida, Monaxonida and Keratosa.
Tetractinellida. These are siliceous demosponges with tetraxon megascleres or with no
skeleton at all. They are rounded, oval, cushion-like or flattened and encrusting. They are
usually unbranched and attach to the substrate either directly or via root spicules. Some
lie free on the bottom. The surface is bristly or hard and stony, or smooth and leathery. An
oscular fringe or rim may be present. The largest tetractinellid is Synops neptuni, which is
cup-shaped and up to 40 cm high. All are marine and occur especially in shallow waters
and along shores.
Most tetractinellids have a thick cortex, though in some there is only a thin dermal layer.
There may be large spaces, the subcortical crypts, between the cortex and choanosome.
The cortex may possess spicules that may project from the surface and may also contain a
fibrous network of fibre cells (desmacytes) parallel to the surface. There may also be a
gelatinous collenchyma layer, containing amoebocytes, outer to the fibrous layer.
In those tetractinellids with a thick cortex, dermal pores open into a canal called a
chone, which may have a sphincter along its length or at the inner end. The dermal pores
may occur in clusters termed pore areas (cribriporal condition) that open into a
subdermal space or a chone. The osculum may have a sieve membrane and oscula may
occur in groups. There is usually no spongocoel. In the absence of a cortex the dermal

pores open almost directly into the flagellated chambers and when the cortex is thin they
open into subdermal spaces.
Chondrosia is an example of a tetractinellid that has no skeleton, and has stiff mesogloea.
Other tetractinellids have skeletons of siliceous spicules, of which the typical type is the
calthrops, which is a tetraxon with 4 equal rays. If the rays are branched then the spicule
is of the candelabra type. One or two rays may be lost. Loss of one ray results in a
triaene with a very long shaft (rhabdome) and small cladi. The cladi curve downward in
the anatriaene, point up in the protriaene and are more or less horizontal in the
plagiotrianes and orthotriaenes. Dichotriaenes have forked cladi. Monaxons (mostly
oxeas) are also present and aster microscleres. The spicules are usually arranged in
radiating bundles and the cladi and the monaxons may project from the surface.
The Tetractinellida is divided into three orders: Myxospongida, Carnosa and Choristida.
Examples of the Myxospongida are Oscarella, Halisarca, Hexidella and Bajulus. The
Carnosa possess calthrops, and examples are Plakina, Plakortis, Thrombus, Corticium,
Corticella, Halina, Dercitopsis, Pachastrella and Chondrilla. Chondrilla has only
spheraster spicules that pack the cortex. The Choristida are divided into the Astrophora
that possess astrose microscleres, e.g. Ancorina, Thenea, Penares, Myriastra, Stellata and
Geodia, and the Sigmatophora that possess sigmas, e.g. Craniella.
Tetractinellids develop through a rhagon-like stage and there is no inversion of the germ
layers. Some have a lithistid skeleton composed of desmas cemented into a framework
and also possess triaenes, astrose and monaxon microscleres, e.g. Corallistes, Kaliapsis,
Pleroma, Theonella, Vetulina and Discodermia.
Monaxonida. The monaxonids have siliceous monaxon megascleres and are the most
common sponges. They occur on the shore down to 6000m. They attach to the substrate
mostly by spongin secretion and form rounded masses, or are bushy or branching, club-,
funnel-, cup- or fan-shaped. There are usually many oscula, which are on the ends of the
branches in branched forms. Cladorhiza is a deep-water monaxonid with a pointed base
encircled by projections that prevent sinkage into the mud. In Polymastia, the incurrent
and excurrent openings are borne on tubes (fistulas) that project from the rounded body.
The monaxonids include the freshwater sponges.
The monaxonids may be radiate (O. Hadromerina) with a thick cortex containing radial
bundles of monaxon megascleres, or reticulate with spongin connecting the spicules into
a network. (Plumose (axinellid) is a variant of the radiate type). Others, e.g.
Halichondria, have no definite spicule arrangement. The order Hadromerina has separate
megascleres and no spongin. In reticulate types the spicules form spongin-coated
networks. The spongin may enclose the spicules (coring) or may partially embed them,
allowing them to project toward the surface (echinating spicules). Some have lithistid
skeletons. The cortex is fibrous and may also be gelatinous. All monaxonids are
leuconoid (eurypylous, aphodal or diplodal).

There are 4 or 5 orders of monaxonid, the Hadromerina, Halichondrina, Poecilosclerina,

Haplosclerina (and the Epipolasida). The Hadromerina include Suberites, Spirastrella,
Cliona, Tethya, Polymastia and Poterion. The Halichondrina includes Halichondria. The
Poecilosclerina includes most demosponges, e.g. Microciona, Myxilla, Tedania, Mycale,
Paresperella, Cladorhiza and Esperiopsis. The Haplosclerina include Haliclona, Reniera,
Chalina and Pachychalina and also the freshwater sponges, e.g. Spongilla, Myenia and
Heteromyenia. The freshwater sponges have zoochlorellae housed inside their
archaeocytes. The order Epipolasida includes Tethya, which is sometimes included in the
Hadromerina.
Keratosa (Horny Sponges). The keratosans possess spongin skeletons and have no
spicules, but they may incorporate rock grains, etc. They are rounded, but the genus
Phyllospongia is leaf-like. The surface is leathery, smooth or covered with small bumps
(conules) that contain the ends of large spongin fibres. Most are black in colour and live
in shallow water attached to hard substrates by spongin secretion. Hircinia contains
filaments of an unknown nature and function (?). The spongin forms a network, either a
lattice or a dendritic network. Some horny sponges have polyaxon spongin spicules.
Some have no skeleton, e.g. Hexidella and Bajulus.
The horny sponge surface is a leathery spongin membrane. Most keratosans are diplodal
leuconoids, but some are eurypylous. The group includes the bath sponges, F. Spongiidae,
e.g. (Eu)spongia and Hippospongia. Bath sponges are thought to live up to 50 years (?).
Other examples of horny sponges are Aplysilla and Verongia.