PHYLUM ECHINODERMATA

The phylum Echinodermata consists of several types of complex organisms which show a general pentameral
symmetry and have a well-developed water vascular system. Echinoderms are also characterized by their
mesodermal skeleton: the skeleton differs from that found in all other invertebrates in that it is secreted by the
middle rather than the outer body layer; thus the test is enveloped in soft tissue. Echinoderms occur in a variety of
morphologies including free-living forms such as starfish and sand dollars as well stalked forms such as sea lilies
which are attached to the sea floor.
The stalked echinoderms also called pelmatozoans are exclusively marine and live in a variety of habitats of
normal salinity. They are all filter-feeders. As a group, the pelmatozoans have been quite abundant in the geologic
past especially the Paleozoic, yet since the close of the Mesozoic they have mainly relegated to deep-water, cryptic
environments.
The non-stalked echinoderms also called Eleutherozoa belongs to the subphyla Asterozoa and Echinozoa, and
are collectively referred to as free-living echinoderms. Similar to the pelmatozoans, the free-living echinoderms all
have a mesodermal skeleton comprised of calcite plates and a complex water-vascular system including tube-feed.
Unlike the pelmatozoans, the groups are quite abundant in the modern seas, occupying a large number
environments by an equally large number of life-habits.

Pelmatozoan general morphology

In general, the pelmatozoan skeleton can be dived into two main parts: The stem and the calyx. The stalk or stem is
composed of numerous disks called columnals which have a central hole called the lumen. Stems are often secured
to the substrate by means of a holdfast or root system. The calyx (sometimes called theca), a cup-like structure,
which may or may not support a variety of arms. The calyx is usually composed of a number of different kinds of
plates, grooves, and pores; some of which are quite specialized

Pelmatozoans Appendage and Calyx Morphology

Ambulacral groove: one of the 5 radially arranged regions specialized for food gathering.
Brachial plates: arm plates; several different types depending on position relative to arm branch e.g.,
primibrachials and secundibrachials.
Brachioles: small erect food gathering appendages surrounding the edge of the ambulacrual area in blastoids.
Pinnules: small linear hair-like branches that may occur on each arm plate and also aid in food gathering.
Basal plates: the circlet of plates below and off-set to, and joining, the radial plates.
Infrabasal plates: secondary plates below basals.
Radial plates: below lowest brachial and above basal plates where ray terminates.
Interray plates: plates that are added between rays.
Lancet: ambulacral plate of blastoid, usually site of brachiole attachment.
Deltoid: small triangular or rhombahedral plate above the radial plates in blastoids.
Pores: openings for tube feet.
Tegmen: Part of calyx occasionally with plates that resides above the attachment points of arms. Sometimes
elevated into a anal pyramid.
Ray: trace of plates from arm through calyx; there are usually five (or multiples thereof) rays in pelmatozoans; ray
terminates with radial plates.

Free-Living (Eleutherozoans) Echinoderm Morphological Terms

The terms below are pertinent to most all of free-living echinoderms and even to stalked pelmatozoans.
Ambulacra: one of the five rays of an echinoderm which support the tube feet; in echinoids it is usually covered
with numerous ambulacral plates.
Interambulacra: region between ambulacral regions.
Apical system: the circlet of 10 plates surrounding the anal system.
Madreporite: a modified opening which commonly has several openings which serve to regulate water pressure.
Periproct: anal opening, usually on the aboral (anti oral) side.
Aristotle's lantern: the mouth, jaw, and teeth parts of an echinoid made up of 40 different skeletal elements.
Spine: rod or club-like structure used in locomotion and/or defense.
Tubercle: attachment structure (usually a knob) to receive a spine.

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1. Classification
Phylum Echinodermata is divided into 5 major groups (Subphylum) that contain 20 Classes, and about half of
which are known only from the Paleozoic (Table 1). The current classification is based on multiple characters such
as general morphology, ossicle structure, arrangement of the water vascular system, and embryology.

TABLE 1. CLASSIFICATION, GENERA, SPECIES & AGE RANGES

1.1 Class Asteroidea (Sea Stars; Starfishes)

The star-shaped echinoderms (Figure 1), the Asteroidea (Sea stars or Starfishes), is the largest, most speciose
(*1600 extant species), diverse and the common class within Phylum Echinodermata and prime predators within
many marine ecosystem. Their size varies from a centimeter to a meter across. They thrive in the intertidal zone,
although, they also occur at depths as great as 10,000 m. The carnivorous asteroids are very sluggish movers and
crawl by the concerted actions of their podia. The Asteroids generally have hollow arms and into this the coelomic
cavity extends. The radial canals are located on the skeleton’s exterior. Their skeleton is rarely robust and consists
of a series of small elements embedded in a collagenous membrane. Hence, after death, they disarticulate quickly,
leaving a poor fossil record. The asteroids first appeared in Early Ordovician but were never common or abundant
in the fossil record, since then. They experienced major faunal transitions concurrently with two large extinction
events; one in Late Devonian and the other in Late Permian.

1.2 Class Ophiuroidea (the Brittle Stars)

The Ophiuroids (Figure 2) are a large group (over 1600 species) that include brittle stars (Ophiurida) and basket
stars (Euryalida). The Ophiuroids are remarkable in that they are able to release their arms at ossicle sutures to
escape predation, hence, they are also called brittle stars. This process of releasing a limb is called Autotomy; the
lost limb is eventually regenerated. The Ophiuroids are somewhat different from the Asteroids (Table 2), the sea
stars, from which they are often mistaken, and hence, often lumped with them in higher taxa, Stelleroidea or
Asterozoa. Unlike asteroids, the ophiuroids are active crawlers with thin whip-like five arms that wriggle like
snakes, hence, their name (ophi = snake, in Greek); they are sometimes also called Serpent stars. The Ophiuroids
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are recorded from the Early Ordovician and achieved global distribution in the Paleozoic. They survived the
Permian–Triassic extinction and increased in numbers during the Mesozoic–Cenozoic.

TABLE 2. COMPARATIVE ACCOUNT OF OPHIUROIDS AND ASTEROIDS

1.3 Classes Holothuroidea and Concentricycloidea (Sea Cucumbers and Sea Daisies)
These soft-bodied sea cucumbers (*200 genera) are without arms. They are the most recently discovered class of
echinoderms; sea daisies are tiny, primitive echinoderms that live at great depths. Sea cucumbers resemble
cucumbers. They possess Sclerites, microscopic hard parts that occur in various shapes resembling hooks, wheels
and anchors. Although, the sea cucumber possess pentaradial symmetry, the anus is opposite to the mouth on an
elongated oral–aboral axis. The Holothurians are the most diverse of the five extant classes of echinoderm, with
over 2000 extant species. However, like the asteroids and the ophiuroids, they also have left a very poor fossil
record and after death, their skeleton is reduced to thousands of microscopic spicules. The only fossilized part are
the 10 ossicles that surrounds the mouth and provide an anchorage for the oral tentacles; forming the Circumoral
ring. The earliest holothurian body fossil is recorded from Late Silurian, however, spicules attributable to
holothurians are known from Ordovician onwards.

1.4 Edrioasteroids
These discoidal, clavate, or subglobular echinoderms (Figure 3) are an extinct group of sessile stem group
eleutherozoans. They are characterized by possessing five ambulacra radiating from a central mouth. The
Stromatocystitids, the earliest forms were completely plated and not attached to the substrate. They represent the
basal eleutherozoans, ancestral to all later forms. The Stromatocystitids first appeared in Early Cambrian, while
isorophids (a derived clade and specialized as hard ground colonizers) appeared a little later, in Late Cambrian, and
survived through to Late Carboniferous.

1.5 Class Crinoidea (Sea Lilies and Feather Stars; Crinoids)

The Crinoids (Figure 4) are unusual looking animals as they look more like plants than animals. Hence their name
“sea lilies”. Superficially, the stem or column of a crinoid resembles the stalk of a flower, the calyx or head
resembles the sepals of a flower, and the arms resemble the petals of a flower. The Crinoids (with over 1000
known genera), are the only echinoderms that live attached to the sea bottom for most of their lives. Two groups of
living crinoids are noted—those with columns, the living stalked crinoids, and those without, the Comatulids.
Typically, a crinoid has a long stem with “roots” or a holdfast (an attachment device) at the bottom, and a cup-
shaped thecum at the top. Some crinoids have become mobile by losing stem; living crinoids are swimmers, and
not attached. Recent crinoids have both a wide vertical distribution (from bathyal depths to tropical reef and
shallow cave habitats) and spatial (from polar to tropical latitudes). The Paleozoic forms were very common on
shallow carbonate platforms. Except in the Cambrian, crinoids were common fossils in the Paleozoic. The normal
arrangement of plates consist of brachials, radials, and basals. They can be dicyclic (having infrabasals) as in the
example or monocyclic (without infrabasals). Crinoids can be divided into 4 subclasses: the Inadunates,
Camerates, Flexibles, and Articulates, only the articulates survive to the Recent.

1.6 Class Blastoidea (Blastoids)

Blastoids are the best known stalked echinoderm of the Paleozoic and widely distributed in Early and Middle
Paleozoic rocks with 95 genera, known, so far. The Extinct Blastoids (such as Pentremites, Figure 5) are
characterized by an armless bud-like calyx on a stem. The primitive stemmed blastoids have pentaradial symmetry
with a very regular distribution of thecal plates; there are 13 thecal plates arranged in three circlets. The theca is the
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size of a rosebud, and a commonly preserved. Although they show pentaradiality, however, diverse body forms,
symmetries, and ambulacral architectures are noted in this group. Blastoids do not have a tegmen. The hydrospire,
a respiratory device, is a distinctive structure in all blastoids hat hangs into the body cavity beneath each
ambulacrum. The blastoids resemble crinoids in appearance and lifestyle. The blastoids first appeared in Middle
Ordovician, peaked in abundance by Mississippian (Early Carboniferous), with a general decline in the Paleozoic
and the final extinction in the Permian. The differences between each class is based primarily upon the pore and
plate arrangement

1.7 Eocrinoids
These stalked arm-bearing echinoderms are a paraphyletic assemblage of basal pelmatozoans with irregularly
plated theca. The eocrinoids include the ancestors of all other blastozoan groups and probably of the crinoids also.
The eocrinoids were the likely ancestors to Diploporita, Rhombifera, Coronoidea, Blastoidea, Parablastoidea and
Paracrinoidea. The Diploporites are characterized by larger and regular plating with no ambulacral flooring plates;
the brachioles arise directly from the thecal plates. The thecal plates are pierced by numerous pairs of pores
(diplopores) that had a respiratory role. The Rhombiferans, arising directly from around the mouth, had stout arms.
The most regular thecal plating is noted in the pentameral symmetry bearing Blastoids that possess three basals,
five radials and five lancet plates. The well-developed ambulacra form an essential part of the theca, giving rise to
a dense fan of brachioles. The oldest blastozoans are eocrinoids of Early Cambrian age (Figure 6). Diploporites
and rhombiferans appeared at beginning of the Ordovician, while the blastoids in the Silurian. The Eocrinoids
finally went extinct by end Permian.

1.8 Class Paracrinoidea (Allied to Crinoids)

The Paracrinoid plates show no definite arrangement but commonly contain pores that are covered (as noted in
deeply weathered specimens). In Camarocystites, the best know genus, the plates of the calyx are concave. The
two arms bear side branchlets which are relatively thick. The Paracrinoid are recorded in Ordovician and Silurian
sediments. They are abundant in mid-Ordovician strata. Some workers believe that Paracrinoid are highly modified
blastozoans.

1.9 Carpoids
The Carpoids (Stylophorans; Class Ctenocystoidea; are unusual forms of the Cambrian strata, and common in the
fossil record from 500 to 300 million years ago. Three competing hypotheses are proposed for the group’s
evolutionary origins:
 Carpoids are very primitive echinoderms with a mobile stalk or single arm filled with muscle
 Stylophorans are advanced echinoderms related to crinoids having an ambulacrum with a tube feet and an
oral tegmen with pharynx, and
 Carpoids are neither of the above two, but primitive chordates that retained a calcite exoskeleton from an
older common ancestor of echinoderms and chordates, with the stalk possessing muscle, notochord and
brain.
Recent studies, based on Middle Cambrian ceratocystid stylophoran from Morocco supports the first
hypothesis.
1.10. Class Echinoidea (sea urchins and sand dollars)
Of the five classes of echinoderm, echinoids are the most diverse and well represented, from shallow
waters to abyssal depths. The Echinoids are exclusively marine benthic macroinvertebrates. These “spiny
skin” forms have an endoskeleton made of hard calcium-rich plates (of single calcite crystal) just beneath
their thin skin. Paleontologically, echinoids are, by far the most significant group. They range in size
from just a few mm in diameter to over 350 mm (both size spectrum is noted in living sea cucumbers and
brittle stars). They also occur in varied shapes (globular, heart-shaped, cylindrical, hemispherical, or even
flattened discoidal). However, irrespective of their overall shape, the skeleton (test) is always constructed
along the same standardized plan. In Echinoids, a rigid and robust calcitic test (skeleton) built by a
mosaic of plates is firmly bound together, thus, making the skeleton is architecturally complex and
mesodermal. Their excellent preservational history (good fossil record) is a testament of the rigid
skeleton and hence, the echinoids are extensively studied for their phylogeny. Contextually, they are also
a valuable palaeobiological tool. Like the crinoid calyx, they are composed of a mosaic of plates; unlike the
crinoids, they are covered by spines. These are tiny bristles in some, but can be quite large in others. The spines

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generally detach from the main skeleton after death, because they are held to it only by non-mineralized ligaments.
There are no arms, and the ambulacral areas (where the perforated plates are found) stretch from pole to pole.
Echinoids can be further divided into two groups: Regular (Figure 7) and Irregular (Figure 8). The extant
echinoid species are equally divided between regular forms (radial, fivefold symmetry; Fig. 9) whose
anus opens in the aboral plated surface (i.e., the anus is located opposite to the mouth), and irregular
forms (bilateral symmetry) whose anus is displaced away from the aboral plates into the posterior
interambulacral zone (Fig. 10). Table 3 lists major differences between Regular and Irregular echinoids.

TABLE 3 MAJOR DIFFERENCES BETWEEN REGULAR AND IRREGULAR ECHINOIDS

REGULAR ECHINOIDS IRREGULAR ECHINOIDS
1 Radially symmetrical Bilaterally symmetrical
2 Anus located in apical system, which is Anus may shift posteriorly out of the apical
large and in centre of aboral surface system, which may be small and not in centre of
aboral surface
3 Five genitals/five oculars in apical system There may be a reduced number of oculars or
genitals
4 Mouth in centre of oral surface Mouth may shift anteriorly
5 Corona usually with large tubercles Corona may have only small granules on surface
6 It has jaws Jaws may be absent
7 Benthonic, mostly epifaunal mobile grazers Benthonic, occur primarily in infaunal
that sometimes occur in rocky subtidal and environments
intertidal environments
8 Have no front or back end and can move in Have a definite front and back and do move in a
any direction particular direction
9 Prefer high energy conditions Prefer low energy conditions
1 Use their tube feet for respiration, to attach Use their tube feet for respiration, digging their
0 themselves to the sea bed and also to move burrows and maintaining them.
1 Ambulacrum runs from center of aboral to Ambulacrum is Modified over test, fused together
1 oral
1 Fasicles are Absent Two present to direct water to mouth and direct
2 waste away
1 Test shape is Circular Test shape distinctively elongate in adult stage
3

Similarities
 Both live in a marine environment
 Both have 5 ambulacrum
 Tube feet used for respiration
 Both have anus and mouth

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