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Ho l o t h u r o i d e a
Sea cucumbers
Al e x a nde r M. Ke r r
t o l - r e v i e w e d
Hol ot hur oi de a
page cont ent
ar t icles & not es
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Explor e Ot her Gr oups
ot her Echinoder mat a
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Tree modified from Kerr (2000).
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Containing group: Echinodermata
I nt r oduct i on
The Holothuroidea, or sea cucumbers, are an abundant and diverse group of worm-like and
usually soft-bodied echinoderms. They are found in nearly every marine environment, but
are most diverse on tropical shallow-water coral reefs. They range from the intertidal, where
they may be exposed briefly at low tide, to the floor of the deepest oceanic trenches. The
oldest undoubted fossils of sea cucumbers are of isolated spicules from the Silurian (ca. 400
million years ago; Gilliland, 1993). Considerable diversification has occurred since then with
about 1400 living species in a variety of forms. Some of these are about 20 cm in length,
though adults of some diminutive species may not exceed a centimeter, while one large
species can reach lengths of 5 m (Synapta maculata). Several species can swim and there
are even forms that live their entire lives as plankton, floating with the ocean currents.
Economically, sea cucumbers are important in two main ways. First, some species produce
toxins that are of interest to pharmaceutical firms seeking to learn their medical value.
Some compounds isolated to date exhibit antimicrobial activity or act as anti-inflammatory
agents and anticoagulants. Second, as a gourmet food item in the orient, they form the
basis of a multimillion-dollar industry that processes the body wall for sale as beche-de-mer
or trepang. However, the high value of some species, the ease with which such shallow-
water forms can be collected and their top-heavy age structures all contribute to over-
exploitation and collapse of the fisheries in some regions. Fishermen in the Pacific islands
use the toxins, some of which act as respiratory inhibitors, to entice fish and octopus from
crevices so that they may be more easily speared. Furthermore, the sticky Cuvierian
tubules (see description below) are placed over bleeding wounds as a bandage.
Char act e r i st i cs
The most important feature distinguishing the sea cucumbers is a calcareous ring that
encircles the pharynx or throat. This ring serves as an attachment point for muscles
operating the oral tentacles and for the anterior ends of other muscles that contract the
body longitudinally. Sea cucumbers are also distinct as echinoderms in having a circlet of
oral tentacles. These may be simple, digitate (with finger-like projections), pinnate (feather-
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like), or peltate (flattened and shield-like). A third key feature, found in 90% of living
species, is the reduction of the skeleton to microscopic ossicles (Figure 1). In some species,
the ossicles may be enlarged and plate-like.
Figure 1. Calcareous skeletal ossicles from the body wall (in situ) of two recent
holothurians.
Left: Wheels and hook-shaped rods of Trochodota allani (Apodida: Chiridotidae).
Right: Spinose wheels with perforated hub and simple rods of Siniotrochus phoxus
(Apodida: Myriotrochidae).
Photographs copyright 2000 Mike Reich.
As in other echinoderms, the holothurian water vascular system consists of an anterior ring
canal from which arise long canals running posteriorly (not shown in Figure 2). Despite their
similarity to the radial canals of other echinoderms, these latter structures arise
embryologically in a quite different manner. For this reason these canals in holothurians
have been recently renamed longitudinal canals (Mooi and David 1997). In holothurians, the
larval structures that would form the radial canals in other echinoderms instead become the
five primary tentacles. Also, holothurians with the exception of members in Elasipodida have
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a madrepore that opens into the coelom (body cavity). In contrast, elasipodans and nearly
all other echinoderms have a madrepore that opens externally.
Figure 2: Main internal anatomical features of a cucumariid sea cucumber
(Dendrochirotida).
Drawing by Ivy Livingstone. Copyright 1995 BIODIDAC.
Some sea cucumbers possess organs not found in other invertebrates. In some
Aspidochirotida, the respiratory trees display Cuvierian tubules. In most species, these are
apparently defensive structures. They can be expelled through the anus, whereupon they
dramatically expand in length and become sticky, entangling or deterring would-be
predators, such as crabs and gastropods. Many forms, with the exception of members of
Elasipodida and Apodida, possess respiratory trees used in gas exchange. These are paired,
heavily branched tubes attached to the intestine near the anus. This type of breathing
("cloacal breathing") is also present in an unrelated group, the echiuran worms.
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Hyman (1955) provides a useful account of holothuroid gross anatomy, Smiley (1994)
covers microscopic aspects, while Smiley et al. (1991) reviews reproduction and larval
development.
The Or de r s of Hol ot hur oi de a
The ancestors of the Apodida, Elasipodida and the lineage leading to the remaining orders
diverged in the middle to late Paleozoic Era between about 350 to 250 million years ago. The
Aspidochirotida, Molpadiida, Dendrochirotida and Dactylochirotida began diverging
somewhat later in the Triassic and Jurassic of the early Mesozoic Era about 200 million years
ago. Assignment to different orders is largely based on the form of the calcareous ring and
tentacles, as well as the presence of certain organs, such as the respiratory trees or the
muscles that retract the oral region.
Descriptions of each order given below are modified from Pawson (1982) and Smiley
(1994):
Apodida
Contains about 269 species in 32 genera and three families. Tentacles are digitate,
pinnate or, in some small species, simple. Respiratory trees are absent. Tube feet
are completely absent. The calcareous ring is without posterior projections. The body
wall is very thin and often transparent. Found in both shallow and deep water.
Elasipodida
Contains about 141 species in 24 genera and five families. Tentacles are shield-
shaped and used in shovelling sediment. Respiratory trees are present. The
calcareous ring is without posterior projections. With the exception of Deimatidae, the
body wall is soft to gelatinous. All forms live in deep water.
Aspidochirotida
There are about 340 species in 35 genera and three families. Tentacles are shield-
shaped. Respiratory trees are present. The calcareous ring is without posterior
projections. The body wall is generally soft and pliant. Most forms live in shallow
water, though one family is restricted to the deep sea.
Molpadiida
There are about 95 species in 11 genera and four families. Tentacles are simple.
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There are about 95 species in 11 genera and four families. Tentacles are simple.
Respiratory trees are present. The calcareous ring is without posterior projections.
The body wall is generally soft and pliant. Most forms live in shallow water, though
one family is restricted to the deep sea.
Dendrochirotida
Contains about 550 species in 90 genera and seven families. Tentacles are highly
branched and extended to filter material from the water column. Respiratory trees
are present. Some members with a calcareous ring composed of numerous small
pieces or having long posterior extensions. Possess muscles for retracting the oral
introvert. The body wall may be hardened from enlarged plate-like ossicles. Live
either attached to hard bottoms or burrow in soft sediment. Most species live in
shallow water.
Dactylochirotida
Contains about 35 species in seven genera and three families. Tentacles are simple
or with a few small digits. Respiratory trees are present. The calcareous ring is
without posterior projections. Possess muscles for retracting the oral introvert. All
members have a rigid body encased in enlarged flattened ossicles. The body is
usually "U" shaped. All members live burrowed in soft sediment. Most live in deep
water.
Di scussi on of Phyl oge ne t i c Re l at i onshi ps
The evolutionary relationships of the major holothuroid lineages were, until quite recently,
poorly understood. This was in part due to their lack of an integrated skeleton like that
providing the extensive fossil record and numerous morphological characters of other
groups of echinoderms. There have been numerous speculations about the relationships
within Holothuroidea extending well back into the 19th century. The methods of modern
comparative biology had not been applied to these problems until quite recently. Then
Littlewood et al. (1997), in an effort to resolve class-level relationships within echinoderms,
sequenced two ribosomal genes from a total of four orders. Their analyses consistently
supported a close relationship between Dendrochirotida and Aspidochirotida, but they could
not resolve the phylogenetic position of Elasipodida and Apodida (Figure 3: A, B). Smith
(1997) subsequently argued that the Elasipodida are more closely related to
(Dendrochirotida + Aspidochirotida) than the Apodida (Figure 3: C). This hypothesis recalls
an early speculation (Semper 1868) whereby Apodida is sister to the remaining
holothuroids.
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Figure 3. Recent hypotheses about holothuroid relationships.
A. Tree based on complete 18S rDNA sequences (Littlewood et al., 1997).
B. Tree based on partial 28S rDNA sequences (Littlewood et al., 1997).
C. Interpretation of the 18S and 28S rDNA data favored by Smith (1997).
More comprehensive cladistic analyses of morphological and DNA data (Kerr, 2000) agree
with Smith's hypothesis. Further, it appears that Dendrochirotida is paraphyletic due to the
Dactylochirotida lineage arising from within the Dendrochirotida. This arrangement of the
two orders is so far supported by few characters, and an alternate arrangement may
ultimately prevail. Kerr (2000) also places Molpadiida as sister to Dendrochirotida plus
Dactylochirotida. Together with Aspidochirotida, the aforementioned orders form a group
united, most notably, by the presence of respiratory trees. The placement of two rare
families currently referred to the Molpadiida, Eupyrgidae and Gephyrothuriidae, is uncertain;
they may turn out to be only distantly related to one another and other ordinal level groups
of holothurians. Based on the presence of respiratory trees, however, they are unlikely to
be closely related to either the Apodida or Elasipodida, which lack such structures. The
remaining features of the higher level relationships shown in the figure at the top of this
page, though, appear solidly supported and unlikely to change with the consideration of new
characters.
Fossi l Hi st or y
As for most soft-bodied animals, holothuroids have a poor fossil record. Published accounts
exist of body fossils for about 19 species, though at least that many body-fossil species lay
undescribed on museum shelves. Most ancient holothuroids are known from fossils of
isolated ossicles. This complicates the taxonomy somewhat since ossicles can differ even
within an individual depending on age, habitat and geography. How then does one identify a
single species? As a result, most fossil holothuroids have been described as paraspecies
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based on unique ossicle types. Entire or isolated elements of the calcareous ring are also
known, though less work has been done on these potentially informative structures. The
rarity of holothuroid fossils in part may be due to a lack of collecting effort, since the
microscopic ossicles require special collecting methods, and there are few specialists
working on the group. In addition, isolated ring elements may sometimes be confused with
the robust plates of other echinoderms.
Figure 4. Isolated pieces of the calcareous rings of fossil holothurians.
Left: Interradial pieces; Center: Radial pieces; both from apodid holothurians from
the Upper Liassic of Germany, approx. 180 Ma;
Right: Pieces from fossil molpadiid holothurians from the Hauterivian of Germany,
approx. 130 Ma.
Photographs copyright 2000 Mike Reich.
Holothuroids probably evolved by at least the Lower Silurian, most likely from a little-known
group of extinct Palaeozoic echinoderms called ophiocistioids. However, the oldest reported
body fossil of a holothuroid is from the Lower Devonian, while the oldest undoubted ossicle
is from the Upper Silurian. Plate ossicles are known from the Ordovician, but their identity as
holothuroid is uncertain because they resemble the plates of other echinoderms. Still, plate
ossicles ascribable to holothuroids are well known and, when combined with the phylogenetic
evidence, suggest that several groups of ancient plated forms existed that are only distantly
related to living plated dendrochirotes and dactylochirotes. Alternatively, these living forms
have retained their ancient armour and Holothuroidea has had a long and repeated history
of losing a plated morphology.
A comprehensive account of holothurian palaeontology is found in Gilliland (1993), while an
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up-to-date bibliography and other palaeontological information is available from Mike Reich's
Fossil Holothuroidea Page.
Ot he r Name s f or Hol ot hur oi de a
holothuroids
Sea cucumbers
Re f e r e nce s
Gilliland, P. M. 1993. The skeletal morphology, systematics and evolutionary history of
holothurians. Special Papers in Palaeontology 47: 1-147
Hyman, L. H. 1955. The Invertebrates. Vol. 4. Echinodermata. New York: McGraw Hill.
Kerr, A. 2000. Evolution and Systematics of Holothuroidea (Echinodermata). Thesis, Yale
University.
Littlewood, D. T. J., A. B. Smith, K. A. Clough and R. H. Emson. 1997. The interrelationships of
the echinoderm classes: morphological and molecular evidence. Biological Journal of the
Linnean Society 61: 409-438.
Mooi, R. and B. David. 1997. Skeletal homologies of echinoderms. Paleontological Society
Papers 3: 305-355.
Pawson, D. L. 1982. Holothuroidea. In: Parker, S. P., ed. Synopsis and Classification of Living
Organisms. New York: McGraw-Hill, 813-818.
Semper, C. 1868. Reisen im Archipel der Philippinen. 2. Wissenschaftliche Resultate. 1.
Holothurien. Leipzig: Wilhelm Engelmann.
Smiley, S., F. S. McEuen, S. Chaffee, and S. Krishnan. 1991. Echinodermata: Holothuroidea.
In: Giese, A. C., J. S. Pearse, and V. B. Pearse, eds. Reproduction of Marine Invertebrates.
Volume 6. Pacific Grove, California: Boxwood Press, 663-750.
Smiley, S. 1994. Holothuroidea. In: Harrison, F. W. and F.-S. Chia, eds. Microscopic Anatomy
of Invertebrates. Volume 14. Echinodermata. New York: Wiley-Liss, 401-471.
Smith, A. B. 1997. Echinoderm larvae and phylogeny. Annual Review of Ecology and
Systematics 28: 219-241.
I nf or mat i on on t he I nt e r ne t
UCMP Berkeley Holothuroidea.
Beche-de-mer Information Bulletin. Coordinated by Chantal Conand.
Taxonomy of Sea Cucumbers. Philip Lambert, Royal British Columbia Museum.
Title Illustrations
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Title Illustrations
Scientific Name Synaptula (Apodida)
Location coral reef on the Eastern Coast
of Thailand
Specimen Condition Live Specimen
Copyright 2000 Sumaitt Putchakarn
Scientific Name Holothuria (Halodeima) atra
(Aspidochirotida)
Location coral reef on the Eastern Coast
of Thailand
Specimen Condition Live Specimen
Copyright 2000 Sumaitt Putchakarn
Scientific Name Cucumaria (Dendrochirotida)
Location Ross Sea, Antarctica
Specimen Condition Live Specimen
Copyright 2000 Norbert Wu
About This Page
Alexander M. Kerr
University of Guam, Mangilao, Guam, USA
Page copyri ght 2000 Al exander M. Kerr
Page: Tree of Life Holothuroidea. Sea cucumbers. Authored by Alexander
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Fi rst onl i ne 01 December 2000
Ci t i ng t hi s page :
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