Sponge

Sponges are animals of the phylum Porifera (/prfr/; meaning "pore bearer"). They are multicellular
organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of
jelly-like mesohyl sandwiched between two thin layers of cells. Sponges have unspecialized cells that can
transform into other types and that often migrate between the main cell layers and the mesohyl in the process.
Sponges do not have nervous, digestive or circulatory systems. Instead, most rely on maintaining a constant water
flow through their bodies to obtain food and oxygen and to remove wastes.

Overview
Sponges are similar to other animals in that they are multicellular, heterotrophic, lack cell walls and produce
sperm cells. Unlike other animals, they lack true tissues and organs, and have no body symmetry. The shapes of
their bodies are adapted for maximal efficiency of water flow through the central cavity, where it deposits
nutrients, and leaves through a hole called the osculum. Many sponges have internal skeletons of spongin and/or
spicules of calcium carbonate or silicon dioxide. All sponges are sessile aquatic animals. Although there are
freshwater species, the great majority are marine (salt water) species, ranging from tidal zones to depths
exceeding 8,800 m (5.5 mi).
While most of the approximately 5,00010,000 known species feed on bacteria and other food particles in the
water, some host photosynthesizing micro-organisms as endosymbionts and these alliances often produce more
food and oxygen than they consume. A few species of sponge that live in food-poor environments have become
carnivores that prey mainly on small crustaceans.[1]
Most species use sexual reproduction, releasing sperm cells into the water to fertilize ova that in some species are
released and in others are retained by the "mother". The fertilized eggs form larvae which swim off in search of
places to settle.[2] Sponges are known for regenerating from fragments that are broken off, although this only
works if the fragments include the right types of cells. A few species reproduce by budding. When conditions
deteriorate, for example as temperatures drop, many freshwater species and a few marine ones produce
gemmules, "survival pods" of unspecialized cells that remain dormant until conditions improve and then either
form completely new sponges or recolonize the skeletons of their parents.[3]
The mesohyl functions as an endoskeleton in most sponges, and is the only skeleton in soft sponges that encrust
hard surfaces such as rocks. More commonly, the mesohyl is stiffened by mineral spicules, by spongin fibers or
both. Demosponges use spongin, and in many species, silica spicules and in some species, calcium carbonate
exoskeletons. Demosponges constitute about 90% of all known sponge species, including all freshwater ones, and
have the widest range of habitats. Calcareous sponges, which have calcium carbonate spicules and, in some
species, calcium carbonate exoskeletons, are restricted to relatively shallow marine waters where production of
calcium carbonate is easiest.[4] The fragile glass sponges, with "scaffolding" of silica spicules, are restricted to
polar regions and the ocean depths where predators are rare. Fossils of all of these types have been found in rocks
dated from 580 million years ago. In addition Archaeocyathids, whose fossils are common in rocks from
530 to 490 million years ago, are now regarded as a type of sponge.
The single-celled choanoflagellates resemble the choanocyte cells of sponges which are used to drive their water
flow systems and capture most of their food. This along with phylogenetic studies of ribosomal molecules have
been used as morphological evidence to suggest sponges are the sister group to the rest of animals. [5] Some
studies have shown that sponges do not form a monophyletic group, in other words do not include all and only
the descendants of a common ancestor. Recent phylogenetic analyses suggest that comb jellies rather than
sponges are the sister group to the rest of animals.[6][7][8][9]
The few species of demosponge that have entirely soft fibrous skeletons with no hard elements have been used by
humans over thousands of years for several purposes, including as padding and as cleaning tools. By the 1950s,
though, these had been overfished so heavily that the industry almost collapsed, and most sponge-like materials
are now synthetic. Sponges and their microscopic endosymbionts are now being researched as possible sources of
medicines for treating a wide range of diseases. Dolphins have been observed using sponges as tools while
foraging.[10]

Distinguishing features
Further information: Cnidaria and Ctenophore
Sponges constitute the phylum Porifera, and have been defined as sessile metazoans (multicelled immobile
animals) that have water intake and outlet openings connected by chambers lined with choanocytes, cells with
whip-like flagella.[11] However, a few carnivorous sponges have lost these water flow systems and the
choanocytes.[12][13] All known living sponges can remold their bodies, as most types of their cells can move within
their bodies and a few can change from one type to another.[13][14]

Like cnidarians (jellyfish, etc.) and ctenophores (comb jellies), and unlike all other known metazoans, sponges'
bodies consist of a non-living jelly-like mass sandwiched between two main layers of cells. [15][16] Cnidarians and
ctenophores have simple nervous systems, and their cell layers are bound by internal connections and by being
mounted on a basement membrane (thin fibrous mat, also known as "basal lamina").[16] Sponges have no nervous
systems, their middle jelly-like layers have large and varied populations of cells, and some types of cells in their
outer layers may move into the middle layer and change their functions.[14]
Sponges[14][15]
Cnidarians and ctenophores[16]
Nervous system
No
Yes, simple
No, except that Homoscleromorpha have Yes: inter-cell connections;
Cells in each layer bound together
basement membranes.[17]
basement membranes
Number of cells in middle "jelly"
Many
Few
layer
Cells in outer layers can move
Yes
No
inwards and change functions

Basic structure
Cell types
A sponge's body is hollow and is held in shape by the mesohyl, a jelly-like substance made mainly of collagen
and reinforced by a dense network of fibers also made of collagen. The inner surface is covered with
choanocytes, cells with cylindrical or conical collars surrounding one flagellum per choanocyte. The wave-like
motion of the whip-like flagella drives water through the sponge's body. All sponges have ostia, channels leading
to the interior through the mesohyl, and in most sponges these are controlled by tube-like porocytes that form
closable inlet valves. Pinacocytes, plate-like cells, form a single-layered external skin over all other parts of the
mesohyl that are not covered by choanocytes, and the pinacocytes also digest food particles that are too large to
enter the ostia,[14][15] while those at the base of the animal are responsible for anchoring it.[15]
Other types of cell live and move within the mesohyl:[14][15]
Lophocytes are amoeba-like cells that move slowly through the mesohyl and secrete collagen fibres.
Collencytes are another type of collagen-producing cell.
Rhabdiferous cells secrete polysaccharides that also form part of the mesohyl.
Oocytes and spermatocytes are reproductive cells.
Sclerocytes secrete the mineralized spicules ("little spines") that form the skeletons of many sponges and
in some species provide some defense against predators.
In addition to or instead of sclerocytes, demosponges have spongocytes that secrete a form of collagen
that polymerizes into spongin, a thick fibrous material that stiffens the mesohyl.
Myocytes ("muscle cells") conduct signals and cause parts of the animal to contract.
"Grey cells" act as sponges' equivalent of an immune system.
Archaeocytes (or amoebocytes) are amoeba-like cells that are totipotent, in other words each is capable of
transformation into any other type of cell. They also have important roles in feeding and in clearing debris
that block the ostia.

Glass sponges' syncytia

Glass sponges present a distinctive variation on this basic plan. Their spicules, which are made of silica, form a
scaffolding-like framework between whose rods the living tissue is suspended like a cobweb that contains most
of the cell types.[14] This tissue is a syncytium that in some ways behaves like many cells that share a single
external membrane, and in others like a single cell with multiple nuclei. The mesohyl is absent or minimal. The
syncytium's cytoplasm, the soupy fluid that fills the interiors of cells, is organized into "rivers" that transport
nuclei, organelles ("organs" within cells) and other substances.[20] Instead of choanocytes, they have further
syncytia, known as choanosyncytia, which form bell-shaped chambers where water enters via perforations. The
insides of these chambers are lined with "collar bodies", each consisting of a collar and flagellum but without a
nucleus of its own. The motion of the flagella sucks water through passages in the "cobweb" and expels it via the
open ends of the bell-shaped chambers.[14]
Some types of cells have a single nucleus and membrane each, but are connected to other single-nucleus cells and
to the main syncytium by "bridges" made of cytoplasm. The sclerocytes that build spicules have multiple nuclei,
and in glass sponge larvae they are connected to other tissues by cytoplasm bridges; such connections between
sclerocytes have not so far been found in adults, but this may simply reflect the difficulty of investigating such
small-scale features. The bridges are controlled by "plugged junctions" that apparently permit some substances to
pass while blocking others.[20]

Water flow and body structures

Most sponges work rather like chimneys: they take in water at the bottom and eject it from the osculum ("little
mouth") at the top. Since ambient currents are faster at the top, the suction effect that they produce by Bernoulli's
principle does some of the work for free. Sponges can control the water flow by various combinations of wholly
or partially closing the osculum and ostia (the intake pores) and varying the beat of the flagella, and may shut it
down if there is a lot of sand or silt in the water.[14]
Although the layers of pinacocytes and choanocytes resemble the epithelia of more complex animals, they are not
bound tightly by cell-to-cell connections or a basal lamina (thin fibrous sheet underneath). The flexibility of these
layers and re-modeling of the mesohyl by lophocytes allow the animals to adjust their shapes throughout their
lives to take maximum advantage of local water currents.[22]
The simplest body structure in sponges is a tube or vase shape known as "asconoid", but this severely limits the
size of the animal. The body structure is characterized by a stalk-like spongocoel surrounded by a single layer of
choanocytes. If it is simply scaled up, the ratio of its volume to surface area increases, because surface increases
as the square of length or width while volume increases proportionally to the cube. The amount of tissue that
needs food and oxygen is determined by the volume, but the pumping capacity that supplies food and oxygen
depends on the area covered by choanocytes. Asconoid sponges seldom exceed 1 mm (0.039 in) in diameter.[14]
Some sponges overcome this limitation by adopting the "syconoid" structure, in which the body wall is pleated.
The inner pockets of the pleats are lined with choanocytes, which connect to the outer pockets of the pleats by
ostia. This increase in the number of choanocytes and hence in pumping capacity enables syconoid sponges to
grow up to a few centimeters in diameter.
The "leuconoid" pattern boosts pumping capacity further by filling the interior almost completely with mesohyl
that contains a network of chambers lined with choanocytes and connected to each other and to the water intakes
and outlet by tubes. Leuconid sponges grow to over 1 m (3.3 ft) in diameter, and the fact that growth in any
direction increases the number of choanocyte chambers enables them to take a wider range of forms, for example
"encrusting" sponges whose shapes follow those of the surfaces to which they attach. All freshwater and most
shallow-water marine sponges have leuconid bodies. The networks of water passages in glass sponges are similar
to the leuconid structure.[14] In all three types of structure the cross-section area of the choanocyte-lined regions is
much greater than that of the intake and outlet channels. This makes the flow slower near the choanocytes and
thus makes it easier for them to trap food particles.[14] For example, in Leuconia, a small leuconoid sponge about
10 centimetres (3.9 in) tall and 1 centimetre (0.39 in) in diameter, water enters each of more than 80,000 intake
canals at 6 cm per minute. However, because Leuconia has more than 2 million flagellated chambers whose
combined diameter is much greater than that of the canals, water flow through chambers slows to 3.6 cm per
hour, making it easy for choanocytes to capture food. All the water is expelled through a single osculum at about
8.5 cm per second, fast enough to carry waste products some distance away.[23]

Skeleton
In zoology a skeleton is any fairly rigid structure of an animal, irrespective of whether it has joints and
irrespective of whether it is biomineralized. The mesohyl functions as an endoskeleton in most sponges, and is
the only skeleton in soft sponges that encrust hard surfaces such as rocks. More commonly the mesohyl is
stiffened by mineral spicules, by spongin fibers or both. Spicules may be made of silica or calcium carbonate, and
vary in shape from simple rods to three-dimensional "stars" with up to six rays. Spicules are produced by
sclerocyte cells,[14] and may be separate, connected by joints, or fused.[13]
Some sponges also secrete exoskeletons that lie completely outside their organic components. For example,
sclerosponges ("hard sponges") have massive calcium carbonate exoskeletons over which the organic matter
forms a thin layer with choanocyte chambers in pits in the mineral. These exoskeletons are secreted by the
pinacocytes that form the animals' skins.[14]

Classes
Sponges were traditionally distributed in three classes: calcareous sponges (Calcarea), glass sponges
(Hexactinellida) and demosponges (Demospongiae). However, studies have shown that the Homoscleromorpha, a
group thought to belong to the Demospongiae, is actually phylogenetically well separated. Therefore, they have
recently been recognized as the fourth class of sponges.[24][25]
Sponges are divided into classes mainly according to the composition of their skeletons:[15]
Spongin Massive
Type of cells[15] Spicules[15]
Body form[15]
fibers[15] exoskeleton[26]
Calcarea
Single
nucleus, Calcite
Never
Common.
Asconoid, syconoid,
single
external May
be
Made of calcite if leuconoid
or
membrane
individual
or
present.
solenoid[27]

large masses
Silica
Mostly syncytia in May
Hexactinellida
all species
individual
fused
Single
nucleus,
Demospongiae
single
external Silica
membrane
Single
nucleus,
Homoscleromorpha single
external Silica
membrane

be
Never
or

Never

Leuconoid

In some species.
In
many
Made of aragonite if Leuconoid
species
present.[13][26]
In
many
Never
species

Sylleibid
leuconoid

or

Vital functions
Movement
Although adult sponges are fundamentally sessile animals, some marine and freshwater species can move across
the sea bed at speeds of 14 mm (0.0390.157 in) per day, as a result of amoeba-like movements of pinacocytes
and other cells. A few species can contract their whole bodies, and many can close their oscula and ostia.
Juveniles drift or swim freely, while adults are stationary.[14]

Respiration, feeding and excretion

Sponges do not have distinct circulatory, respiratory, digestive, and excretory systems instead the water flow
system supports all these functions. They filter food particles out of the water flowing through them. Particles
larger than 50 micrometers cannot enter the ostia and pinacocytes consume them by phagocytosis (engulfing and
internal digestion). Particles from 0.5 m to 50 m are trapped in the ostia, which taper from the outer to inner
ends. These particles are consumed by pinacocytes or by archaeocytes which partially extrude themselves
through the walls of the ostia. Bacteria-sized particles, below 0.5 micrometers, pass through the ostia and are
caught and consumed by choanocytes.[14] Since the smallest particles are by far the most common, choanocytes
typically capture 80% of a sponge's food supply.[26] Archaeocytes transport food packaged in vesicles from cells
that directly digest food to those that do not. At least one species of sponge has internal fibers that function as
tracks for use by nutrient-carrying archaeocytes,[14] and these tracks also move inert objects.[15]
It used to be claimed that glass sponges could live on nutrients dissolved in sea water and were very averse to silt.
[28]
However a study in 2007 found no evidence of this and concluded that they extract bacteria and other microorganisms from water very efficiently (about 79%) and process suspended sediment grains to extract such prey.[29]
Collar bodies digest food and distribute it wrapped in vesicles that are transported by dynein "motor" molecules
along bundles of microtubules that run throughout the syncytium.[14]
Sponges' cells absorb oxygen by diffusion from water into cells as water flows through body, into which carbon
dioxide and other soluble waste products such as ammonia also diffuse. Archeocytes remove mineral particles
that threaten to block the ostia, transport them through the mesohyl and generally dump them into the outgoing
water current, although some species incorporate them into their skeletons.[14]

Carnivorous sponges
A few species that live in waters where the supply of food particles is very poor prey on crustaceans and other
small animals. So far only 137 species have been discovered. [30] Most belong to the family Cladorhizidae, but a
few members of the Guitarridae and Esperiopsidae are also carnivores.[31] In most cases little is known about how
they actually capture prey, although some species are thought to use either sticky threads or hooked spicules.[31][32]
Most carnivorous sponges live in deep waters, up to 8,840 m (5.49 mi),[33] and the development of deep-ocean
exploration techniques is expected to lead to the discovery of several more. [14][31] However one species has been
found in Mediterranean caves at depths of 1723 m (5675 ft), alongside the more usual filter feeding sponges.
The cave-dwelling predators capture crustaceans under 1 mm (0.039 in) long by entangling them with fine
threads, digest them by enveloping them with further threads over the course of a few days, and then return to
their normal shape; there is no evidence that they use venom.[33]
Most known carnivorous sponges have completely lost the water flow system and choanocytes. However the
genus Chondrocladia uses a highly modified water flow system to inflate balloon-like structures that are used for
capturing prey.[31][34]

Endosymbionts
Freshwater sponges often host green algae as endosymbionts within archaeocytes and other cells, and benefit
from nutrients produced by the algae. Many marine species host other photosynthesizing organisms, most
commonly cyanobacteria but in some cases dinoflagellates. Symbiotic cyanobacteria may form a third of the total
mass of living tissue in some sponges, and some sponges gain 48% to 80% of their energy supply from these

micro-organisms.[14] In 2008 a University of Stuttgart team reported that spicules made of silica conduct light into
the mesohyl, where the photosynthesizing endosymbionts live.[35] Sponges that host photosynthesizing organisms
are most common in waters with relatively poor supplies of food particles, and often have leafy shapes that
maximize the amount of sunlight they collect.[15]
A recently discovered carnivorous sponge that lives near hydrothermal vents hosts methane-eating bacteria, and
digests some of them.[15]

"Immune" system
Sponges do not have the complex immune systems of most other animals. However they reject grafts from other
species but accept them from other members of their own species. In a few marine species, gray cells play the
leading role in rejection of foreign material. When invaded, they produce a chemical that stops movement of
other cells in the affected area, thus preventing the intruder from using the sponge's internal transport systems. If
the intrusion persists, the grey cells concentrate in the area and release toxins that kill all cells in the area. The
"immune" system can stay in this activated state for up to three weeks.[15]

Reproduction
Asexual
Sponges have three asexual methods of reproduction: after fragmentation; by budding; and by producing
gemmules. Fragments of sponges may be detached by currents or waves. They use the mobility of their
pinacocytes and choanocytes and reshaping of the mesohyl to re-attach themselves to a suitable surface and then
rebuild themselves as small but functional sponges over the course of several days. The same capabilities enable
sponges that have been squeezed through a fine cloth to regenerate. [36] A sponge fragment can only regenerate if it
contains both collencytes to produce mesohyl and archeocytes to produce all the other cell types. [26] A very few
species reproduce by budding.[37]
Gemmules are "survival pods" which a few marine sponges and many freshwater species produce by the
thousands when dying and which some, mainly freshwater species, regularly produce in autumn. Spongocytes
make gemmules by wrapping shells of spongin, often reinforced with spicules, round clusters of archeocytes that
are full of nutrients.[38] Freshwater gemmules may also include phytosynthesizing symbionts. [39] The gemmules
then become dormant, and in this state can survive cold, drying out, lack of oxygen and extreme variations in
salinity.[14] Freshwater gemmules often do not revive until the temperature drops, stays cold for a few months and
then reaches a near-"normal" level.[39] When a gemmule germinates, the archeocytes round the outside of the
cluster transform into pinacocytes, a membrane over a pore in the shell bursts, the cluster of cells slowly emerges,
and most of the remaining archeocytes transform into other cell types needed to make a functioning sponge.
Gemmules from the same species but different individuals can join forces to form one sponge. [40] Some gemmules
are retained within the parent sponge, and in spring it can be difficult to tell whether an old sponge has revived or
been "recolonized" by its own gemmules.[39]
Sexual
Most sponges are hermaphrodites (function as both sexes simultaneously), although sponges have no gonads
(reproductive organs). Sperm are produced by choanocytes or entire choanocyte chambers that sink into the
mesohyl and form spermatic cysts while eggs are formed by transformation of archeocytes, or of choanocytes in
some species. Each egg generally acquires a yolk by consuming "nurse cells". During spawning, sperm burst out
of their cysts and are expelled via the osculum. If they contact another sponge of the same species, the water flow
carries them to choanocytes that engulf them but, instead of digesting them, metamorphose to an ameboid form
and carry the sperm through the mesohyl to eggs, which in most cases engulf the carrier and its cargo.[41]
A few species release fertilized eggs into the water, but most retain the eggs until they hatch. There are four types
of larvae, but all are balls of cells with an outer layer of cells whose flagellae or cilia enable the larvae to move.
After swimming for a few days the larvae sink and crawl until they find a place to settle. Most of the cells
transform into archeocytes and then into the types appropriate for their locations in a miniature adult sponge.[41]
Glass sponge embryos start by dividing into separate cells, but once 32 cells have formed they rapidly transform
into larvae that externally are ovoid with a band of cilia round the middle that they use for movement, but
internally have the typical glass sponge structure of spicules with a cobweb-like main syncitium draped around
and between them and choanosyncytia with multiple collar bodies in the center. The larvae then leave their
parents' bodies.[42]
Life cycle
Sponges in temperate regions live for at most a few years, but some tropical species and perhaps some deepocean ones may live for 200 years or more. Some calcified demosponges grow by only 0.2 mm (0.0079 in) per
year and, if that rate is constant, specimens 1 m (3.3 ft) wide must be about 5,000 years old. Some sponges start
sexual reproduction when only a few weeks old, while others wait until they are several years old.[14]

Coordination of activities

Adult sponges lack neurons or any other kind of nervous tissue. However most species have the ability to
perform movements that are coordinated all over their bodies, mainly contractions of the pinacocytes, squeezing
the water channels and thus expelling excess sediment and other substances that may cause blockages. Some
species can contract the osculum independently of the rest of the body. Sponges may also contract in order to
reduce the area that is vulnerable to attack by predators. In cases where two sponges are fused, for example if
there is a large but still unseparated bud, these contraction waves slowly become coordinated in both of the
"Siamese twins". The coordinating mechanism is unknown, but may involve chemicals similar to
neurotransmitters.[43] However glass sponges rapidly transmit electrical impulses through all parts of the
syncytium, and use this to halt the motion of their flagella if the incoming water contains toxins or excessive
sediment.[14] Myocytes are thought to be responsible for closing the osculum and for transmitting signals between
different parts of the body.[15]
Sponges contain genes very similar to those that contain the "recipe" for the post-synaptic density, an important
signal-receiving structure in the neurons of all other animals. However, in sponges these genes are only activated
in "flask cells" that appear only in larvae and may provide some sensory capability while the larvae are
swimming. This raises questions about whether flask cells represent the predecessors of true neurons or are
evidence that sponges' ancestors had true neurons but lost them as they adapted to a sessile lifestyle.[44]

Ecology
Habitats
Sponges are worldwide in their distribution, living in a wide range of ocean habitats, from the polar regions to the
tropics.[26] Most live in quiet, clear waters, because sediment stirred up by waves or currents would block their
pores, making it difficult for them to feed and breathe. [28] The greatest numbers of sponges are usually found on
firm surfaces such as rocks, but some sponges can attach themselves to soft sediment by means of a root-like
base.[45]
Sponges are more abundant but less diverse in temperate waters than in tropical waters, possibly because
organisms that prey on sponges are more abundant in tropical waters.[46] Glass sponges are the most common in
polar waters and in the depths of temperate and tropical seas, as their very porous construction enables them to
extract food from these resource-poor waters with the minimum of effort. Demosponges and calcareous sponges
are abundant and diverse in shallower non-polar waters.[47]
The different classes of sponge live in different ranges of habitat:
Type
of
Water type[15]
Depth[15]
surface[15]
Calcarea
Marine
less than 100 m (330 ft)
Hard
Soft or firm
Glass sponges Marine
Deep
sediment
[15]
Inter-tidal to abyssal;
a carnivorous
Marine, brackish; and about
Demosponges
demosponge has been found at 8,840 m (5.49 mi) Any
150 freshwater species[14]
[33]

As primary producers
Sponges with photosynthesizing endosymbionts produce up to three times more oxygen than they consume, as
well as more organic matter than they consume. Such contributions to their habitats' resources are significant
along Australia's Great Barrier Reef but relatively minor in the Caribbean.[26]

Defenses
Close-up of the sponge boring Entobia in a modern oyster valve. Note the chambers which are connected by
short tunnels.
Many sponges shed Sponge spicules, forming a dense carpet several meters deep that keeps away echinoderms
which would otherwise prey on the sponges.[26] They also produce toxins that prevent other sessile organisms
such as bryozoans or sea squirts from growing on or near them, making sponges very effective competitors for
living space. One of many examples includes ageliferin.
A few species, the Caribbean fire sponge Tedania ignis, cause a severe rash in humans who handle them. [14]
Turtles and some fish feed mainly on sponges. It is often said that sponges produce chemical defenses against
such predators.[14] However an experiment showed that there is no relationship between the toxicity of chemicals
produced by sponges and how they taste to fish, which would diminish the usefulness of chemical defenses as
deterrents. Predation by fish may even help to spread sponges by detaching fragments.[15]
Glass sponges produce no toxic chemicals, and live in very deep water where predators are rare.[28]

Predation

Sponge flies, also known as spongilla-flies (Neuroptera, Sisyridae), are specialist predators of freshwater
sponges. The female lays her eggs on vegetation overhanging water. The larvae hatch and drop into the water
where they seek out sponges to feed on. They use their elongated mouthparts to pierce the sponge and suck the
fluids within. The larvae of some species cling to the surface of the sponge while others take refuge in the
sponge's internal cavities. The fully grown larvae leave the water and spin a cocoon in which to pupate.[48]

Bioerosion
The Caribbean chicken-liver sponge Chondrilla nucula secretes toxins that kill coral polyps, allowing the
sponges to grow over the coral skeletons. [14] Others, especially in the family Clionaidae, use corrosive substances
secreted by their archeocytes to tunnel into rocks, corals and the shells of dead mollusks.[14] Sponges may remove
up to 1 m (3.3 ft) per year from reefs, creating visible notches just below low-tide level.[26]

Diseases
Caribbean sponges of the genus Aplysina suffer from Aplysina red band syndrome. This causes Aplysina to
develop one or more rust-colored bands, sometimes with adjacent bands of necrotic tissue. These lesions may
completely encircle branches of the sponge. The disease appears to be contagious and impacts approximately 10
percent of A. cauliformis on Bahamian reefs.[49] The rust-colored bands are caused by a cyanobacterium, but it is
unknown whether this organism actually causes the disease.[49][50]

Collaboration with other organisms

In addition to hosting photosynthesizing endosymbionts,[14] sponges are noted for their wide range of
collaborations with other organisms. The relatively large encrusting sponge Lissodendoryx colombiensis is most
common on rocky surfaces, but has extended its range into seagrass meadows by letting itself be surrounded or
overgrown by seagrass sponges, which are distasteful to the local starfish and therefore protect Lissodendoryx
against them; in return the seagrass sponges get higher positions away from the sea-floor sediment.[51]
Shrimps of the genus Synalpheus form colonies in sponges, and each shrimp species inhabits a different sponge
species, making Synalpheus one of the most diverse crustacean genera. Specifically, Synalpheus regalis utilizes
the sponge not only as a food source, but also as a defense against other shrimp and predators. [52] As many as
16,000 individuals inhabit a single loggerhead sponge, feeding off the larger particles that collect on the sponge
as it filters the ocean to feed itself.[53]

Evolutionary history
24-isopropylcholestane is a stable derivative of 24-isopropylcholesterol, which is said to be produced by
demosponges but not by eumetazoans ("true animals", i.e. cnidarians and bilaterians). Since choanoflagellates are
thought to be animals' closest single-celled relatives, a team of scientists examined the biochemistry and genes of
one choanoflagellate species. They concluded that this species could not produce 24-isopropylcholesterol but that
investigation of a wider range of choanoflagellates would be necessary in order to prove that the fossil 24isopropylcholestane could only have been produced by demosponges.[54] Although a previous publication reported
traces of the chemical 24-isopropylcholestane in ancient rocks dating to 1,800 million years ago,[55] recent
research using a much more accurately dated rock series has revealed that these biomarkers only appear before
the end of the Marinoan glaciation approximately 635 million years ago,[56] and that "Biomarker analysis has yet
to reveal any convincing evidence for ancient sponges pre-dating the first globally extensive Neoproterozoic
glacial episode (the Sturtian, ~713 million years ago in Oman)". Nevertheless, this 'sponge biomarker' could have
other sources such as marine algae so may not constrain the origin of Porifera.[57]
Although molecular clocks and biomarkers suggest sponges existed well before the Cambrian explosion of life,
silica spicules like those of demosponges are absent from the fossil record until the Cambrian, [58] although one
unsubstantiated report exists of spicules in rocks dated around 750 million years ago,[59] although this appears
unlikely based on the above reference. Well-preserved fossil sponges from about 580 million years ago in the
Ediacaran period have been found in the Doushantuo Formation. These fossils, which include spicules,
pinacocytes, porocytes, archeocytes, sclerocytes and the internal cavity, have been classified as demosponges.
Fossils of glass sponges have been found from around 540 million years ago in rocks in Australia, China and
Mongolia.[60] Early Cambrian sponges from Mexico belonging to the genus Kiwetinokia show evidence of fusion
of several smaller spicules to form a single large spicule. [61] Calcium carbonate spicules of calcareous sponges
have been found in Early Cambrian rocks from about 530 to 523 million years ago in Australia. Other probable
demosponges have been found in the Early Cambrian Chengjiang fauna, from 525 to 520 million years ago.[62]
Freshwater sponges appear to be much younger, as the earliest known fossils date from the Mid-Eocene period
about 48 to 40 million years ago.[60] Although about 90% of modern sponges are demosponges, fossilized remains
of this type are less common than those of other types because their skeletons are composed of relatively soft
spongin that does not fossilize well.[63] Earliest sponge symbionts are known from the early Silurian.[64]

Archaeocyathids, which some classify as a type of coralline sponge, are common in the Cambrian period from
about 530 million years ago, but apparently died out by the end of the Cambrian 490 million years ago.[62]

Family tree
A choanoflagellate
Fungi
Choanoflagellates
Metazoa

Glass sponges
Demosponges
Calcareous sponges
Eumetazoa

Comb jellies
Placozoa
Cnidaria
(jellyfish, etc.)

Simplified
family
as closest to more complex animals[65]

tree

showing

calcareous

sponges

Plants
Fungi
Metazoa

Most demosponges
Calcareous sponges
Homoscleromorpha
Eumetazoa

Cnidaria
(jellyfish, etc.)
Other metazoans

Simplified
family
as closest to more complex animals[66]

tree

showing

Homoscleromorpha

In the 1990s sponges were widely regarded as a monophyletic group, all of them having descended from a
common ancestor that was itself a sponge, and as the "sister-group" to all other metazoans (multi-celled animals),
which themselves form a monophyletic group. On the other hand, some 1990s analyses also revived the idea that
animals' nearest evolutionary relatives are choanoflagellates, single-celled organisms very similar to sponges'
choanocytes which would imply that most Metazoa evolved from very sponge-like ancestors and therefore that
sponges may not be monophyletic, as the same sponge-like ancestors may have given rise both to modern
sponges and to non-sponge members of Metazoa.[65]
Analyses since 2001 have concluded that Eumetazoa (more complex than sponges) are more closely related to
particular groups of sponges than to the rest of the sponges. Such conclusions imply that sponges are not
monophyletic, because the last common ancestor of all sponges would also be a direct ancestor of the Eumetazoa,
which are not sponges. A study in 2001 based on comparisons of ribosome DNA concluded that the most
fundamental division within sponges was between glass sponges and the rest, and that Eumetazoa are more
closely related to calcareous sponges, those with calcium carbonate spicules, than to other types of sponge.[65] In
2007 one analysis based on comparisons of RNA and another based mainly on comparison of spicules concluded
that demosponges and glass sponges are more closely related to each other than either is to calcareous sponges,
which in turn are more closely related to Eumetazoa.[60][67]

Other anatomical and biochemical evidence links the Eumetazoa with Homoscleromorpha, a sub-group of
demosponges. A comparison in 2007 of nuclear DNA, excluding glass sponges and comb jellies, concluded that:
Homoscleromorpha are most closely related to Eumetazoa; calcareous sponges are the next closest; the other
demosponges are evolutionary "aunts" of these groups; and the chancelloriids, bag-like animals whose fossils are
found in Cambrian rocks, may be sponges.[66] The sperm of Homoscleromorpha share with those of Eumetazoa
features that those of other sponges lack. In both Homoscleromorpha and Eumetazoa layers of cells are bound
together by attachment to a carpet-like basal membrane composed mainly of "type IV" collagen, a form of
collagen not found in other sponges although the spongin fibers that reinforce the mesohyl of all demosponges
is similar to "type IV" collagen.[17]
A comb jelly
The analyses described above concluded that sponges are closest to the ancestors of all Metazoa, of all multicelled animals including both sponges and more complex groups. However, another comparison in 2008 of 150
genes in each of 21 genera, ranging from fungi to humans but including only two species of sponge, suggested
that comb jellies (ctenophora) are the most basal lineage of the Metazoa included in the sample. If this is correct,
either modern comb jellies developed their complex structures independently of other Metazoa, or sponges'
ancestors were more complex and all known sponges are drastically simplified forms. The study recommended
further analyses using a wider range of sponges and other simple Metazoa such as Placozoa.[68] The results of
such an analysis, published in 2009, suggest that a return to the previous view may be warranted. 'Family trees'
constructed using a combination of all available data morphological, developmental and molecular concluded
that the sponges are in fact a monophyletic group, and with the cnidarians form the sister group to the bilaterians.
[69]

Archaeocyathids are very common fossils in rocks from the Early Cambrian about 530 to 520 million years ago
but are not found after the Late Cambrian. It has been suggested that they were produced by: sponges; cnidarians;
algae; foraminiferans; a completely separate phylum of animals, Archaeocyatha; or even a completely separate
kingdom of life, labeled Archaeata or Inferibionta. Since the 1990s archaeocyathids have been regarded as a
distinctive group of sponges.[70]
It is difficult to fit chancelloriids into classifications of sponges or more complex animals. An analysis in 1996
concluded that they were closely related to sponges on the grounds that the detailed structure of chancellorid
sclerites ("armor plates") is similar to that of fibers of spongin, a collagen protein, in modern keratose (horny)
demosponges such as Darwinella.[72] However another analysis in 2002 concluded that chancelloriids are not
sponges and may be intermediate between sponges and more complex animals, among other reasons because
their skins were thicker and more tightly connected than those of sponges. [73] In 2008 a detailed analysis of
chancelloriids' sclerites concluded that they were very similar to those of halkieriids, mobile bilaterian animals
that looked like slugs in chain mail and whose fossils are found in rocks from the very Early Cambrian to the Mid
Cambrian. If this is correct, it would create a dilemma, as it is extremely unlikely that totally unrelated organisms
could have developed such similar sclerites independently, but the huge difference in the structures of their bodies
makes it hard to see how they could be closely related.[71]