Hickman−Roberts−Larson: 6.

Sponges: Phylum Text © The McGraw−Hill

Animal Diversity, Third Porifera Companies, 2002
Edition

chapter
• • • • • •

s i x
Sponges
Phylum Porifera

The Advent of Multicellularity

Sponges are the simplest multicellular animals. Because the cell is the
elementary unit of life, organisms larger than unicellular protozoa
arose as aggregates of such building units. Nature has experimented
with producing larger organisms without cellular differentiation—
certain large, single-celled marine algae, for example—but such exam-
ples are rarities. There are many advantages to multicellularity as
opposed to simply increasing the mass of a single cell. Since it is at
cell surfaces that metabolic exchange takes place, dividing a mass
into smaller units greatly increases the surface area available for meta-
bolic activities. It is impossible to maintain a workable surface-to-mass
ratio by simply increasing the size of a single-celled organism. Thus,
multicellularity is a highly adaptive path toward increasing body size.
Strangely, while sponges are multicellular, their organization is
quite distinct from other metazoans. A sponge body is an assemblage
of cells embedded in a gelatinous matrix and supported by a skele-
ton of minute needlelike spicules and protein. Because sponges nei-
ther look nor behave like other animals, it is understandable that
they were not completely accepted as animals by zoologists until
well into the nineteenth century. Nonetheless, molecular evidence
suggests that sponges share a common ancestor with other metazoa.

A Caribbean demosponge,
Aplysina fistularis.
105
 Hickman−Roberts−Larson: 6. Sponges: Phylum Text © The McGraw−Hill
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Edition

106 chapter six

ponges belong to phylum Porifera (po-rif´-er-a) (L. porus, spicules or spongin [a specialized collagen] or both). A fourth

S pore, + fera, bearing). Sponges bear myriads of tiny

pores and canals that constitute a filter-feeding system
adequate for their inactive life habit. They are sessile animals
class (Sclerospongiae) was created to contain sponges with a
massive calcareous skeleton and siliceous spicules. Some zool-
ogists maintain that known species of sclerosponges can be
and depend on water currents carried through their unique placed in the traditional classes of sponges (Calcarea and Demo-
canal systems to bring them food and oxygen and to carry spongiae); thus, we do not need a new class.
away their body wastes. Their bodies are little more than
masses of cells embedded in a gelatinous matrix and stiffened
by a skeleton of minute spicules of calcium carbonate or sil- Ecological Relationships
ica and collagen (p. 56). They have no organs or true tissues,
and even their cells show a certain degree of independence. As Most of the 5000 or more sponge species are marine, although
sessile animals with only negligible body movement, they have some 150 species live in fresh water. Marine sponges are abun-
not evolved a nervous system or sense organs and have only dant in all seas and at all depths, and a few even exist in brack-
the simplest of contractile elements. ish water.Although their embryos are free swimming,adults are
So, although they are multicellular, sponges share few of always attached, usually to rocks, shells, corals, or other sub-
the characteristics of other metazoan phyla. For this reason merged objects (figure 6.2). Some bottom-dwelling forms even
they are often called Parazoa (Gr. para, beside or alongside of, grow on sand or mud. Their growth patterns often depend on
+ zōon, animal). shape of the substratum, direction and speed of water currents,
Sponges vary in size from a few millimeters to the great and availability of space, so that the same species may differ
loggerhead sponges, which may reach 2 m or more across. markedly in appearance under different environmental condi-
Many sponge species are brightly colored because of pigments tions. Sponges in calm waters may grow taller and straighter
in their dermal cells. Red, yellow, orange, green, and purple than those in rapidly moving waters.
sponges are not uncommon. However, color fades quickly Many animals (crabs, nudibranchs, mites, bryozoans, and
when sponges are removed from water. Some sponges, includ- fish) live as commensals or parasites in or on sponges. Larger
ing the simplest, are radially symmetrical, but many are quite sponges particularly tend to harbor a large variety of inverte-
irregular in shape. Some stand erect, some are branched or brate commensals. On the other hand, sponges grow on many
lobed, and others are low, even encrusting, in form (figure 6.1). other living animals, such as molluscs, barnacles, brachiopods,
Some bore holes into shells or rocks. corals, or hydroids. One sponge has been described that preys
Sponges are an ancient group, with an abundant fossil on shrimp. Some crabs attach pieces of sponge to their cara-
record extending back to the early Cambrian period and even, pace for camouflage and for protection, since most predators
according to some claims, the Precambrian. Living poriferans seem to find sponges distasteful. Certainly one reason for the
traditionally have been assigned to three classes: Calcarea success of sponges as a group is that they have few enemies.
(with calcareous spicules), Hexactinellida (six-rayed siliceous Because of a sponge’s elaborate skeletal framework and often
spicules), and Demospongiae (with a skeleton of siliceous noxious odor, most potential predators find sampling a sponge

f i g u r e 6.1
Some growth habits and forms of sponges.

Finger sponge
Tube sponge

Encrusting sponge
Red boring sponge Variable
sponge

Coral head
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Sponges: Phylum Porifera 107

position in animal kingdom

Multicellular organisms (metazoa) are typically divided Although Mesozoa and Parazoa are multicellular, their
into three grades: (1) Mesozoa (a single phylum), (2) Para- plan of organization is distinct from that in the eumetazoan
zoa (phylum Porifera, sponges; and phylum Placozoa), and phyla. Such cellular layers as they possess are not homolo-
(3) Eumetazoa (all other phyla). Space limitations preclude gous to the germ layers of the Eumetazoa, and neither group
covering the very small phyla Mesozoa and Placozoa in has developmental patterns resembling those of other meta-
this text. zoa. The name Parazoa means the “beside-animals.”

biological contributions
1. Although the simplest in organization of all metazoa, are organized into incipient tissues of a low level of
these groups do compose a higher level of morpholog- integration.
ical and physiological integration than that found in 3. Developmental patterns of poriferans are different
protozoan colonies. Mesozoa and Parazoa may be said from those of other phyla, and their embryonic layers
to belong to a cellular level of organization. are not homologous to the germ layers of Eumetazoa.
2. Sponges (poriferans) have several types of cells 4. Sponges have developed a unique system of water
differentiated for various functions, some of which currents on which they depend for food and oxygen.

water outlet. These openings are connected by a system of

canals, some of which are lined with peculiar flagellated collar
cells called choanocytes, whose flagella maintain a current
of environmental water through the canals. Water enters the
canals through a multitude of tiny incurrent pores (dermal
ostia) and leaves by way of one or more large oscula.
Choanocytes not only keep the water moving but also trap
and phagocytize food particles that are carried in the water.
Cells lining the passageways are very loosely organized. Col-
lapse of the canals is prevented by the skeleton, which,
depending on the species, may be composed of needlelike
calcareous or siliceous spicules, a meshwork of organic spon-
gin fibers, or a combination of the two.
Sessile, or almost sessile, animals make few movements
and therefore need little in the way of nervous, sensory, or
f i g u r e 6.2 locomotor parts. Sponges apparently have been sessile from
This orange demosponge, Mycale laevis, often grows beneath platelike their earliest appearance and have never acquired specialized
colonies of the stony coral, Montastrea annularis. The large oscula of the nervous or sensory structures, and they have only the very sim-
sponge are seen at the edges of the plates. Unlike some other sponges, plest of contractile systems.
Mycale does not burrow into the coral skeleton and may actually protect
the coral from invasion by more destructive species. Pinkish radioles of a
Christmas tree worm, Spirobranchus giganteus (phylum Annelida, class Types of Canal Systems
Polychaeta), also project from the coral colony. An unidentified reddish
sponge can also be seen further to the right of and below the Christmas Most sponges have one of three types of canal systems—
tree worm. asconoid, syconoid, or leuconoid (figure 6.3).

about as pleasant as eating a mouthful of glass splinters

embedded in evil-smelling gristle. Some reef fishes, however, Asconoids—Flagellated Spongocoels
do graze on shallow-water sponges.
Asconoid sponges have the simplest organization. They are
small and tube shaped.Water enters through microscopic der-
Form and Function mal pores into a large cavity called a spongocoel, which is
lined with choanocytes. Choanocyte flagella pull water
The only body openings of these unusual animals are pores, through the pores and expel it through a single large osculum
usually many tiny ones called ostia for incoming water, and (figure 6.3). Leucosolenia (Gr. leukos, white, + solen, pipe) is
one to a few large ones called oscula (sing., osculum) for an asconoid type of sponge. Its slender, tubular individuals
 Hickman−Roberts−Larson: 6. Sponges: Phylum Text © The McGraw−Hill
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108 chapter six

Osculum
Spicule Excurrent canal
Osculum
Flagellated
Pinacocyte Osculum Radial canal chamber
Choanocyte
Ostium Incurrent
Spicule canal

Porocyte Dermal Incurrent

ostium canal
Prosopyle

Spongocoel Apopyle

Ostium

Asconoid (Leucosolenia) Syconoid (Sycon) Leuconoid (Euspongia)

f i g u r e 6.3
Three types of canal systems. The degree of complexity from simple asconoid to complex leuconoid type has involved mainly the water canal and
skeletal systems, accompanied by outfolding and branching of the collar-cell layer. The leuconoid type is considered the major plan for sponges because
it permits greater size and more efficient water circulation.

characteristics of phylum
porifera
1. Multicellular; body a loose aggregation of cells of
mesenchymal origin
2. Body with pores (ostia), canals, and chambers
that serve for passage of water
3. All aquatic; mostly marine
4. Symmetry radial or none
5. Epidermis of flat pinacocytes; most interior
surfaces lined with flagellated collar cells
(choanocytes) that create water currents; a
gelatinous protein matrix called mesohyl
(mesoglea) contains amebocytes, collencytes, f i g u r e 6.4
and skeletal elements Clathrina canariensis (class Calcarea) is common on Caribbean reefs in
6. Skeletal structure of fibrillar collagen (a protein) caves and under ledges.
and calcareous or siliceous crystalline spicules,
often combined with variously modified collagen
(spongin) fibrils grow in groups attached by a common stolon, or stem, to
7. No organs or true tissues; digestion intracellular; objects in shallow seawater. Clathrina (L. clathri, lattice work)
excretion and respiration by diffusion is an asconoid with bright yellow, intertwined tubes (figure
8. Reactions to stimuli apparently local and indepen- 6.4). Asconoids are found only in the Calcarea.
dent; nervous system probably absent
9. All adults sessile and attached to substratum
Syconoids—Flagellated Canals
10. Asexual reproduction by buds or gemmules and
sexual reproduction by eggs and sperm; free- Syconoid sponges look somewhat like larger editions of
swimming ciliated larvae asconoids, from which they were derived. They have a tubular
body and single osculum, but the body wall, which is thicker
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Sponges: Phylum Porifera 109

Collencyte
Radial canal lined Archaeocyte Pinacocyte
with choanocytes

Prosopyles

Amphiblastula larva

Dermal ostium

Mesohyl

Incurrent
canals
lined with
pinacocytes Mesohyl
Choanocyte Spicules
Spongocoel
f i g u r e 6.6
Apopyle Small section through sponge wall, showing four types of sponge cells.
Pinacocytes are protective and contractile; choanocytes create water
currents and engulf food particles; archaeocytes have a variety of func-
f i g u r e 6.5 tions, including phagocytosis of food particles and differentiation into
Cross section through wall of sponge Sycon, showing canal system. other cell types; collencytes appear to have a contractile function.

and more complex than that of asconoids, receives water The mesohyl is the “connective tissue” of the sponges; in it are
through incurrent canals that deliver it to the choanocyte- found various ameboid cells, fibrils, and skeletal elements. Sev-
lined radial canals, which empty into the spongocoel (fig- eral types of cells occur in sponges.
ures 6.3 and 6.5). The spongocoel in syconoids is lined with
epithelial-type cells, rather than choanocytes as found in
asconoids. Syconoids are found in Calcarea. Sycon (Gr. sykon, a Pinacocytes
fig) is a commonly studied example of the syconoid type of The nearest approach to a true tissue in sponges is arrange-
sponge (figure 6.5). ment of the pinacocyte (figure 6.6) cells of the external
epithelium. These are thin, flat, epithelial-type cells that cover
the exterior surface and some interior surfaces. Some are T-
Leuconoids—Flagellated Chambers
shaped, with their cell bodies extending into the mesohyl.
Leuconoid organization is the most complex of the sponge Pinacocytes are somewhat contractile and help regulate the
types and permits an increase in sponge size. Most leuconoids surface area of the sponge. Some pinacocytes are modified as
form large masses with numerous oscula (see figure 6.2). Clus- contractile myocytes, which are usually arranged in circular
ters of flagellated chambers are filled from incurrent canals and bands around the oscula or pores, where they help regulate
discharge water into excurrent canals that eventually lead to the rate of water flow. Myocytes contain microfilaments similar
the osculum (see figure 6.3). Most sponges are of the leuconoid to those found in muscle cells of other animals.
type, which occurs in most Calcarea and in all other classes.
These three types of canal systems—asconoid, syconoid,
and leuconoid—demonstrate an increase in complexity and Porocytes
efficiency of the water-pumping system, but they do not imply Tubular cells that pierce the wall of asconoid sponges, through
an evolutionary or developmental sequence. The leuconoid which water flows, are called porocytes (see figure 6.3).
grade of construction has evolved independently many times
in sponges. Possession of a leuconoid plan is of clear adaptive
value; it increases the proportion of flagellated surfaces com- Choanocytes
pared with the volume, thus providing more collar cells to Choanocytes, which line flagellated canals and chambers, are
meet food demands. ovoid cells with one end embedded in mesohyl and the other
exposed. The exposed end bears a flagellum surrounded by a
collar (figures 6.6 and 6.7). Electron microscopy shows that
Types of Cells the collar is made up of adjacent microvilli, connected to each
Sponge cells are loosely arranged in a gelatinous matrix called other by delicate microfibrils, so that the collar forms a fine fil-
mesohyl (also called mesoglea, or mesenchyme) (figure 6.6). tering device for straining food particles from the water (figure
 Hickman−Roberts−Larson: 6. Sponges: Phylum Text © The McGraw−Hill
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Edition

110 chapter six

Excurrent canal
f i g u r e 6.7
Food trapping by sponge cells. A, Cutaway section Water flow
of canals showing cellular structure and direction Food route
Pinacocyte
of water flow. B, Two choanocytes. C, Structure
H 2O
of the collar. Small red arrows indicate movement
Apopyle
of food particles.
A Incurrent
Archaeocyte
canal
Choanocyte
Spicule

Collencyte Prosopyle
H 2O
Flagellum

Trapped food Collar microvilli

B particle
Microfibrils
Food vacuole C

Nucleus H 2O
H 2O

H 2O H 2O

6.7B and C). The beat of a flagellum pulls water through the Glass sponges have siliceous spicules with six rays arranged in
sievelike collar and forces it out through the open top of the three planes at right angles to each other. There are many varia-
collar. Particles too large to enter the collar become trapped in tions in the shape of spicules, and these structural variations are
secreted mucus and slide down the collar to the base where of taxonomic importance.
they are phagocytized by the cell body. Larger particles have
already been screened out by the small size of the dermal
pores and prosopyles. Food engulfed by the cells is passed on Sponge Physiology
to a neighboring archaeocyte for digestion.
Sponges feed primarily on particles suspended in water
pumped through their canal systems. Detritus particles, plank-
Archaeocytes tonic organisms, and bacteria are consumed nonselectively in
the size range from 50 µm (average diameter of ostia) to 0.1 µm
Archaeocytes are ameboid cells that move about in the (width of spaces between the microvilli of the choanocyte col-
mesohyl (see figure 6.6) and carry out a number of functions. lar). Pinacocytes may phagocytize particles at the surface, but
They can phagocytize particles at the external epithelium and most larger particles are consumed in the canals by archaeo-
receive particles for digestion from choanocytes. Archaeocytes cytes that move close to the lining of the canals. The smallest
apparently can differentiate into any of the other types of particles, accounting for about 80% of the particulate organic
more specialized cells in the sponge. Some, called sclero- carbon, are phagocytized by choanocytes. Digestion is entirely
cytes, secrete spicules. Others, called spongocytes, secrete intracellular (occurs within cells), a chore performed by the
the spongin fibers of the skeleton, and collencytes secrete fib- archaeocytes.
rillar collagen. Sponges consume a significant portion of their nutrients
in the form of organic matter dissolved in water circulating
through the system. Such material is apparently taken up by
Types of Skeletons pinocytosis (or potocytosis).
Its skeleton gives support to a sponge, preventing collapse of There are no respiratory or excretory organs; these func-
canals and chambers. The major structural protein in the animal tions are performed by diffusion. Contractile vacuoles have
kingdom is collagen, and fibrils of collagen are found through- been found in archaeocytes and choanocytes of freshwater
out the intercellular matrix of all sponges. In addition, various sponges.
Demospongiae secrete a form of collagen traditionally known as All life activities of the sponge depend on the current of
spongin. Demospongiae also secrete siliceous spicules. Calcare- water flowing through the body. A sponge pumps a remarkable
ous sponges secrete spicules composed mostly of crystalline amount of water. Some large sponges can filter 1500 liters of
calcium carbonate that have one, three, or four rays (figure 6.8). water a day. At least some sponges can crawl (move laterally
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Sponges: Phylum Porifera 111

f i g u r e 6.8
A, Types of spicules found in sponges. There
is amazing diversity, beauty, and complexity
of form among the many types of spicules.
B, Some examples of sponge body forms.

A Siliceous spicules Siliceous spicules Spongin Calcareous

(Hexactinellida) (Demospongiae)

Callyspongia

Leucosolenia

B Poterion Euplectella

over their supporting substratum) at speeds of up to 4 mm per Other sponges are oviparous, and both oocytes and
day. This ability may give them an advantage over more sessile sperm are expelled free into the water. The free-swimming
encrusting organisms in competition for space. larva of most sponges is a solid-bodied parenchymula (figure
6.9). The outwardly directed, flagellated cells migrate to the
interior after the larva settles and become the choanocytes in
Reproduction and Development the flagellated chambers. In sexual reproduction ova are fertil-
All sponges are capable of both sexual and asexual reproduc- ized by motile sperm in the mesohyl; there the zygotes
tion. In sexual reproduction most sponges are monoe- develop into flagellated larvae, which break loose and are car-
cious (have both male and female sex cells in one individual). ried away by water currents.
Sperm arise from transformation of choanocytes. In Calcarea The loose organization of sponges is ideally suited for
and at least some Demospongiae, oocytes also develop from regeneration of injured and lost parts, and for asexual repro-
choanocytes; in other demosponges oocytes apparently are duction.Sponges reproduce asexually by forming external buds
derived from archaeocytes. Sperm are released into the water that detach or remain to form colonies. In addition to external
by one individual and are taken into the canal system of buds, which all sponges can form, freshwater sponges and
another. There choanocytes phagocytize them, then transform some marine sponges reproduce asexually by the regular for-
into carrier cells and carry the sperm through the mesohyl to mation of internal buds called gemmules (figure 6.10). These
the oocytes. dormant masses of encapsulated archaeocytes are produced

Parenchymula
Excurrent Incurrent
larva
opening opening

Developing demosponge

f i g u r e 6.9
Development of demosponges.
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112 chapter six

Secondary Primary
Micropyle reticulum reticulum
Inner
membrane

Archaeocytes

Spicules
Trabecular
reticulum
Flagellated
chamber
Choanoblast
f i g u r e 6.10
Section through a gemmule of a freshwater sponge (Spongillidae). Gem- Mesohyl
mules are a mechanism for survival of the harsh conditions of winter.
On return of favorable conditions, the archaeocytes exit through the
micropyle to form a new sponge. The archaeocytes of the gemmule give
H2O
rise to all cell types of the new sponge. Incurrent
space

Collar
during unfavorable conditions. They can survive periods of bodies
drought and freezing and more than three months in the
absence of oxygen. Later, with the return of favorable condi- Nuclei of Prosopyle
trabecular
tions for growth, archaeocytes in the gemmules escape and reticulum
develop into new sponges.
f i g u r e 6.11
Diagram of part of a flagellated chamber of hexactinellids. The primary
Brief Survey of Sponges and secondary reticula are branches of the trabecular reticulum, which
is syncytial. Cell bodies of the choanoblasts and their processes are
borne by the primary reticulum and are embedded in a thin, collagenous
Class Calcarea (Calcispongiae) mesohyl. Processes of the choanoblasts end in collar bodies, whose
Calcarea are calcareous sponges, so called because their collars extend up through the secondary reticulum. Flagellar action
propels water (arrows) to be filtered through the mesh of collar microvilli.
spicules are composed of calcium carbonate. Spicules are
straight monaxons or have three or four rays (see figure 6.8A).
The sponges tend to be small—10 cm or less in height—and reticulum separate into a primary reticulum (incurrent side)
tubular or vase shaped. They may be asconoid, syconoid, or and a secondary reticulum (excurrent, or atrial side) (figure
leuconoid in structure. Although many are drab, some are 6.11). The spherical choanoblasts are borne by the primary
bright yellow, red, green, or lavender. Leucosolenia, Clathrina reticulum, and each choanoblast has one or more processes
(see figure 6.4), and Sycon are common examples. extending to collar bodies, the bases of which are also sup-
ported by the primary reticulum. Each collar and flagellum
extends into the flagellated chamber through an opening in the
Class Hexactinellida (Hyalospongiae) secondary reticulum. Water is drawn into the space between
Glass sponges are nearly all deep-sea forms. Most are radially primary and secondary reticula through prosopyles in the pri-
symmetrical and range from 7 to 10 cm to more than 1 m in mary reticulum, then through the collars into the lumen of the
length. One distinguishing feature, reflected in the class name, flagellated chamber. Collar bodies do not participate in phago-
is the skeleton of six-rayed siliceous spicules bound together cytosis, but rather that process is accomplished by the primary
in an exquisite glasslike latticework (see figure 6.8A). and secondary reticula.
Their tissue structure differs so dramatically from other The latticelike network of spicules found in many glass
sponges that some scientists advocate placing hexactinellids in sponges is of exquisite beauty, such as that of Euplectella
a subphylum separate from other sponges. The body of hexac- (NL. from Gr. euplektos, well-plaited), a classic example of
tinellids is composed of a single, continuous syncytial tissue Hexactinellida.
called a trabecular reticulum. The trabecular reticulum is
the largest, continuous syncytial tissue known in Metazoa. It is
bilayered and encloses a thin, collagenous mesohyl between
Class Demospongiae
the layers, as well as cellular elements such as archaeocytes, Demospongiae comprise approximately 80% of all sponge
sclerocytes, and choanoblasts. Choanoblasts are associated species, including most larger sponges. Their skeletons may be
with flagellated chambers, where the layers of the trabecular of siliceous spicules, spongin fibers, or both. All members of
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Sponges: Phylum Porifera 113

A B C

f i g u r e 6.12
Marine Demospongiae on Caribbean coral reefs. A, Pseudoceratina crassa is a colorful sponge growing at moderate depths. B, Ectyoplasia ferox is
irregular in shape and its oscula form small, volcano-like cones. It is toxic and may cause skin irritation if touched. C, Monanchora unguifera with
commensal brittle star, Ophiothrix suensoni (phylum Echinodermata, class Ophiuroidea).

the class are leuconoid, and all are marine except one family, reefs. The Devonian period saw rapid development of many
the freshwater Spongillidae. Freshwater sponges are widely glass sponges. The possibility that sponges arose from choano-
distributed in well-oxygenated ponds and streams, where they flagellates (protozoa that bear collars and flagella) earned sup-
are found encrusting plant stems and old pieces of submerged port for a time. However, many zoologists object to that
wood. They resemble a bit of wrinkled scum, pitted with hypothesis because sponges do not acquire collars until later
pores, and are brownish or greenish in color. Freshwater in their embryological development. The outer cells of the lar-
sponges die and disintegrate in late autumn, leaving gemmules vae are flagellated but not collared, and they do not become
to survive the winter. collar cells until they become internal. Also, collar cells are
Marine Demospongiae are varied in both color and found in certain corals and echinoderms, so they are not
shape. Some are encrusting; some are tall and fingerlike; and unique to the sponges.
some are shaped like fans, vases, cushions, or balls (figure 6.12). However, these objections are countered by evidence
Some sponges bore into and excavate molluscan shells and based on the sequences of ribosomal RNA. This evidence
coral skeletons. Loggerhead sponges may grow several meters supports the hypothesis of a common ancestor for choano-
in diameter. So-called bath sponges belong to the group called flagellates and metazoans. It suggests also that sponges and
horny sponges, which have only spongin skeletons. They can Eumetazoa are sister groups, with Porifera having split off
be cultured by cutting out pieces of the individual sponges, fas- before the origin of the radiates and placozoans, but sharing
tening them to a weight, and dropping them into the proper a common ancestor.
water conditions. It takes many years for them to grow to mar-
ket size. Most commercial “sponges” now on the market are
synthetic, but the harvest and use of bath sponges persist. Adaptive Radiation
Porifera are a highly successful group that includes several
thousand species and a variety of marine and freshwater habi-
Phylogeny and Adaptive Radiation tats. Their diversification centers largely on their unique water-
current system and its various degrees of complexity.
Proliferation of flagellated chambers in leuconoid sponges was
Phylogeny more favorable to an increase in body size than that of
Sponges originated before the Cambrian period. Two groups asconoid and syconoid sponges because facilities for feeding
of calcareous spongelike organisms occupied early Paleozoic and gaseous exchange were greatly enlarged.
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114 chapter six

classification of phylum porifera

Class Calcarea (cal-ca´re-a) (L. calcis, lime, + Gr. spongos, cal or funnel shaped. Flagellated chambers in simple
sponge) (Calcispongiae). Have spicules of calcium syconoid or leuconoid arrangement. Habitat mostly deep
carbonate that often form a fringe around the osculum; water; all marine. Examples: Venus’ flower basket
spicules needle-shaped or three- or four-rayed; all three (Euplectella), Hyalonema.
types of canal systems (asconoid, syconoid, leuconoid)
Class Demospongiae (de-mo-spun´je-e) (tolerated
represented; all marine. Examples: Sycon, Leucosolenia,
misspelling of Gr. desmos, chain, tie, bond, + spongos,
Clathrina.
sponge). Have skeleton of siliceous spicules that are not
Class Hexactinellida (hex-ak-tin-el´i-da) (Gr. hex, six, + six-rayed, or spongin, or both. Leuconoid-type canal
aktis, ray) (Hyalospongiae). Have six-rayed, siliceous systems. One family found in fresh water; all others marine.
spicules extending at right angles from a central point; Examples: Thenea, Cliona, Spongilla, Myenia, and all bath
spicules often united to form network; body often cylindri- sponges.

summary
Sponges (phylum Porifera) are an abundant gen in the form of large fibers or filaments flagellated cells at the surface to the interior
marine group with some freshwater represen- (spongin),calcareous or siliceous spicules,or a (parenchymella). Sponges have great regener-
tatives.They have various specialized cells,but combination of spicules and spongin in most ative abilities.
these cells are not organized into tissues or species. Sponges are an ancient group, remote
organs. They depend on the flagellar beat of Sponges reproduce asexually by bud- phylogenetically from other metazoa, but mo-
their choanocytes to circulate water through ding, fragmentation, and gemmules (internal lecular evidence suggests that they are the
their bodies for gathering food and exchange buds). Most sponges are monoecious but pro- sister group to Eumetazoa. Their adaptive
of respiratory gases. They are supported by duce sperm and oocytes at different times. radiation is centered on elaboration of the
secreted skeletons of fibrillar collagen, colla- Embryogenesis is unusual, with a migration of water circulation and filter-feeding system.

review questions
1. Give six characteristics of sponges. 7. Describe the skeletons of each of the 12. What are possible ancestors to
2. Briefly describe asconoid, syconoid, classes of sponges. sponges? Justify your answer.
and leuconoid body types in sponges. 8. Describe how sponges feed, respire, 13. It has been suggested that despite
3. What sponge body type is most and excrete. being large, multicellular animals,
efficient and makes possible the 9. What is a gemmule? sponges function more like protozoa.
largest body size? 10. Describe how gametes are produced What aspects of sponge biology
4. Define the following: ostia, osculum, and the process of fertilization in most support this statement and how?
spongocel, mesohyl. sponges. Consider, for example, nutrition,
5. Define the following: pinacocytes, 11. What is the largest class of sponges, reproduction, gas exchange, and
choanocytes, archaeocytes, and what is its body type? cellular organization.
sclerocytes, collencytes.
6. What material is found in the skeleton
of all sponges?
 Hickman−Roberts−Larson: 6. Sponges: Phylum Text © The McGraw−Hill
Animal Diversity, Third Porifera Companies, 2002
Edition

Sponges: Phylum Porifera 115

selected references
See also general references on page 406. Leys, S. P. 1999. The choanosome of Wood, R. 1990. Reef-building sponges. Am.
Bergquist, P. R. 1978. Sponges. Berkeley, Cali- hexactinellid sponges. Invert. Biol. 118: Sci. 78:224–235. The author presents
fornia, University of California Press. 221–235. Choanosomes are flagellated evidence that the known sclerosponges
Excellent monograph on sponge struc- chambers and associated tissues. This belong to either the Calcarea or the
ture, classification, evolution, and gen- author supports the position that Demospongiae and that a separate
eral biology. Hexactinellida should constitute a class Sclerospongiae is not needed.
Bond, C. 1997. Keeping up with the sponges. separate subphylum. She includes an Wyeth, R. C. 1999.Video and electron
Nat. Hist. 106:22–25. Sponges are not excellent description of trabecular microscopy of particle feeding in
fixed in permanent position; they can reticulum. sandwich cultures of the hexactinellid
crawl on their substrate. Haliclona Reiswig, H. M., and T. L. Miller. 1998. Fresh- sponge, Rhabdocalyptus dawsoni.
loosanoffi can move over 4 mm/day. water sponge gemmules survive months Invert. Biol. 118:236–242. Phagocytosis is
Gould, S. J. 1995. Reversing established of anoxia. Invert. Biol. 117:1–8. Hatcha- not by choanoblasts but by trabecular
orders. Nat. Hist. 104(9):12–16. bility of gemmules kept in the absence reticulum, especially primary reticulum.
Describes several anomalous animal of oxygen was equal to controls, but He places Hexactinellida in subphylum
relationships, including the sponge they would not hatch unless oxygen Symplasma and the rest of Porifera in
that preys on shrimp. was present. subphylum Cellularia.

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