I
was recently asked to provide some more information about medusa worms
in one of the forums, and while searching the web to provide some links,
I realized that there is very little information out there about the
biology and aquarium care of these fascinating animals.So, I decided that I would take this opportunity to write a
column about one of my favorite critters to try to provide some
information about these interesting animals.
OK, granted they are
probably not as colorful or attractive as many of the animals that
we find on coral reefs, and to most people they do look pretty
much like a worm.But
ever since I first saw one, back while cruising the local pet shop
in high school, I have always had a soft spot (my wife tells me it
is between my ears) for these animals.Regardless of whether or not they are the prettiest of reef
organisms, I have always been fascinated by Medusa worms!I think that given the appropriate conditions and care, the
unique body form of these animals, combined with the pinnate
feeding tentacles and high level of activity make for an
attractive and popular addition to many reef aquaria.I’ll get to what the appropriate conditions and care are
by the end of the article, but I should start off by explaining
that the name is misleading, as medusa worms are not worms at all,
but rather are legless (Apodid) sea cucumbers.These animals are members of the Family Synaptidae in the
Class Holothuroidea and the Phylum Echinodermata.
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Proud sponsor of this column
Although
medusa worms generally don’t look much like an echinoderm, other
members of the Phylum Echinodermata ought to be very familiar to
most reef aquarists.The
roughly 7-10,000 living species of Echinoderms have a diverse
arrangement of body plans including: sea stars (Asteroids),
brittle stars (Ophiuroids), sea urchins (Echinoids), feather stars
(Crinoids) and the group to which medusa worms belong: the sea
cucumbers (Holothuroids).One
thing that links all these diverse body forms includes a complex
water vascular system that is composed of a series of fluid-filled
canals that allow them to move via hydraulic action.The most obvious components of this hydraulic system to us
as aquarists will be the muscular podia (which are often
sucker-tipped, and better-known as tube feet), that the animals
use to move around.If you’re interested in more information about the general
biology of these groups, I suggest that you pick up a good
invertebrate zoology textbook, such as Brusca and Brucsa (1990) or
Ruppert and Barnes (1994).
Sea
cucumbers are all members of the Class Holothuroidea, but members of
this Class can be surprisingly different from one another.Within the Class Holothuroidea, there are three sub-classes: the
Dendrochirotacea (largely suspension-feeding sea cucumbers such as the
popular sea apple, Pseudocolochirus spp.), the Aspidochirotacea
(largely deposit-feeding sea cucumbers such as the popular tiger-tail
cucumber, Holthuria spp.), and the Apodacea (the unusual group of
legless cucumbers to which the medusa worms belong).Unlike the other Classes of echinoderms mentioned above, in which
the mouth is on the "bottom side" of the animal, the sea
cucumbers all lie on their side, and the mouth is located at the “font
end” of the animal.Although
both ends generally look pretty similar to the casual observer, the sure
way to identify the head is to see the oral tentacles with which the
animal feeds.Depending on
the species of sea cucumber and what they eat, these oral tentacles can
be either stuck out into the water column to suspension feed, or applied
directly to the substrate to deposit feed.The legless synaptid cucumbers fall into this latter group, and
are the most active deposit feeders of the Holothuroids.That tends to make the head end of these animals quite obvious,
because they almost always have their feeding tentacles extended as they
move about in search of particulate detritus to eat.Now although I mentioned above that tube feet are one of the
unifying characteristics of the echinoderms, these Apodid sea cucumbers
are called “legless” because they lack obvious tube feet.In this case, it is because the ancestor of these cucumbers
actually had tube feet, but they have been lost through time in this
group.That is not to say
that they are always absent: some Apodid cukes really do lack any
detectable vestiges of tube feet, but in many species the tube feet are
simply reduced to the point that they are not really visible.
It
is also important to note that aside from their lack of tube feet, most
apodid cucumbers lack the respiratory tree and most of the associated
structures characteristic of the other sub-classes of sea cucumbers.Instead, these animals tend to gain oxygen and expel carbon
dioxide primarily across the body surfaces.Because these animals lack a respiratory tree and the associated
tubules of Cuvier (defensive structures with which the most potent
toxins of sea cucumbers are associated), they are relatively nontoxic in
comparison to some of their more potent relatives such as the sea apples
(Pseudocolochirus spp.) and spotted sea cucumbers (in particular,
Bohadaschia argus and Actinopyga agassizii).The Cuvierian tubules commonly found in species of Holothuria,
Bohadaschia, Stichopus and Actinopyga liberate a highly toxic
saponin compound known as holothurin.This compound acts to quickly stun and even kill potential
predators, which are effectively asphyxiated (suffocated) by the toxin.The toxins of some of these species are so effective that South
Pacific island cultures use the macerated bodies of these sea cucumbers
to stun and capture fishes, crabs and lobsters as a method of
traditional fishing (Frey 1951; Ruppert and Barnes 1994)!Despite the fact that the medusa worms lack these structures and
are considered ‘relatively non-toxic’ by comparison to many of these
species, this does not mean that they are by any means non-toxic.For example, the beautiful sea apples (e.g., Pseudocolochirus
violaceus and P. tricolor) also lack these Cuverian tubules,
but along with Bohadachia argus and Actinopyga agassizii,
are considered among the most toxic sea cucumbers in the world.In fact, according to Wilkens (1998), it takes only about 1g of
tissue from any of these particularly toxic species to poison the fish
in a 25g tank.These toxins
also affect humans, and many people suffer moderate to severe skin and
eye irritation if they come into contact with these toxins (Cunningham
and Goetz 1996).Some cases
in which toxins came into direct contact with the eyes have resulted in
blindness, and deaths have even been reported in cases where people have
eaten these animals without the proper preparation.
In the most general
terms, virtually any soft-bodied animal that would make easy
prey on a coral reef, such as these sea cucumbers, will
typically be defended in some way by distasteful chemicals or
physical armament, and medusa worms are no exception.Like virtually all sea cucumbers, they are soft-bodied,
lacking any real physical armament, and so they all tend to have
a variety of nasty chemicals associated with their bodies to
deter predators from feeding on them.Even without the Cuvierian tubules and their potent
toxins, medusa worms have a variety of distasteful chemicals
associated with the skin and body wall to protect them from
being eaten by fishes, crabs and lobsters on the coral reef.Although the specific toxins associated with synaptid
cucumbers are somewhat different from those of most other
cucumbers studied to date (Kuznetsova et al. 1989; Ponomarenko
et al. 2001), these animals are still reported to be highly
toxic to fishes in marine aquaria if they are seriously injured
(e.g., Delbeek and Sprung 1994; Fenner 2000; Michael undated
online; Sprung 2001).In
general, toxins are only released when the cucumbers are under
severe stress (such as being chewed up after being sucked into a
powerhead or overflow grate), and a diligent aquarist will
usually prevent this from ever happening, and therefore never
experience any problems with one of these animals.If however, and accident happens, and the cucumber is
stressed severely enough to release its chemical defenses, then
a good water change, together with an efficient skimmer and some
activated carbon are usually sufficient to prevent any fish from
asphyxiating from the soap-like holothurin.I will not detour further into the chemical defenses of
sea cucumbers in this article, however I may come back to the
subject another time.
Bohadaschia
spp. live on open sandy rubble zones where it feeds on
organic detritus and benthic microalgae mopped from the
sediments by it's palmate feeding tentacles. It
rarely buries itself in the sand, but occasionally
collects tiny stones and pieces of algae to camouflage
itself. (photo by Julian Sprung)
The
distinctive bright ocelli (false eye spots) covering the
body of this Bohadaschia spp. may serve as warning
coloration to predators that these cucumbers are capable
of expelling sticky defensive threads together highly
toxic chemicals to protect themselves. These
defensive threads, known as Cuverian tubules are
extruded in response to extreme stress, and although
this cucumber is quite attractive, it is also one of the
most toxic species that could quickly wipe out an entire
aquarium if stressed. (Photo: Julian Sprung)
Medusa worms get their
common name because they resemble a giant worm from which a
"mop" of feeding tentacles is constantly being slapped
across the substrate before being drawn into the mouth.These animals are impossible to positively identify by
anyone other than an expert, and even then it usually involves
knowing exactly where the animal was collected, and likely
killing the animal and examining internal structures to be
certain of the species identification.Therefore, chances are that you'll never know which
species have been imported when you see one offered for sale in
the local pet shop.The
most common genera to be imported for the aquarium trade are Euapta,
Synapta, Synaptula or Opheodesoma.All of these animals look fairly similar, being rather
soft and flaccid, with large rounded knobs (sometimes likened to
a string of pearls) along the length of the body.All of these cucumbers seen in the pet trade are typically
shallow water animals (most species are rarely seen below 50ft),
but although they look fairly similar to our eye, they can have
dramatically different biology and care requirements.Although they are not usually an obvious presence on most
coral reefs, apodid sea cucumbers are a common member of
virtually all coral reefs throughout the Caribbean and
Indo-Pacific.Many
species can get quite large (some species can exceed 6 feet in
length!).The
reason that such a large and ubiquitous member of coral reef
communities is only infrequently observed is that they are
almost entirely nocturnal (only active at night).In the aquarium, however, they often lose this strict
nocturnal schedule and are often seen cruising around the tank
in full daylight (I will come back to this issue later in the
article).
Despite the
similarity in appearance, these animals have a wide range of feeding
habits.For example, the
Caribbean Synaptula hydriformis appears to be primarily a
generalist herbivore, feeding primarily on diatoms, with the occasional
piece of red or green algal detritus ingested when they can find it (Martínez
1989), and these animals are common in a variety of habitats including
coral reefs, sea grass beds, mangroves, and even inland salt-water lakes
(Hendler et al. 1995; Pawson 1986)!Other species, such as Synaptula lamperti, are highly
specialized feeders which are only associated with living individuals of
the sponge Ianthella basta (which I have never seen offered for
sale in any pet shop).These
cucumbers live by ingesting tiny organic particles and exudates from the
surface of the sponge, and appear to require these specific sponge
metabolites to defend and nourish it (Hammond and Wilkinson 1989).These two species of Synaptula appear fairly similar, but
obviously it will be much easier to accommodate the first animal in an
aquarium than the second!Other species, such as the Caribbean Euapta lappa, are
generalist detritivores that cruise the sea floor at night collecting
any tiny particles of organic detritus within a given size range from
the reef rubble and base rock structure.Some species appear to graze from a wide range of sediment types
and particle sizes, while others are highly specific (Hammond 1982a).For example, Leptosynapta multigranula, graze organic
detritus from only carbonate sediments of varying sizes, while other
species, such as Leptosynapta tenuis or L.crassipatina appear
to feed effectively only while burrowing through very fine sands and
muddy bottoms (Hendler et al. 1995).In some areas of the South Eastern US, Leptosynapta tenuis
are the major consumers of detritus, and in suitable habitats, Ruppert
and Fox (1988) estimated that these cucumbers can process as much as 25
metric tons of sediment per hectare per year!In contrast, Chiridota rotifera is found actively grazing
detritus from surfaces of tide pools, sea grass beds, calcareous algae,
coral rubble fields, coral heads, and sandy mudflats – not exactly
what anyone would consider a habitat specialist.
The
primary problem with getting any of these animals, however, is that you
cannot identify them by yourself, and will therefore have no idea of
which species you are buying.That leaves you with no option but to trust your supplier about
its care and requirements.That is easier for some people than others, and depending on how
reliable and knowledgeable your supplier is you may be able to obtain a
generalist detritivore that will thrive in a well-established reef
aquarium.If, however, you simply purchase an animal at random, there are
more specific feeders than generalists in this group.Therefore, you have a fair to excellent chance that you will get
a species that is a highly specific feeder, and for which you may have
no way to provide food.For this reason I generally recommend that people avoid buying
one of these animals unless they can first determine what they feed
upon.
Furthermore,
the few studies that have been done on the feeding habits of these
animals indicate that they are very active feeders, and being among the
most active of echinoderms, they require a lot of food energy.Research indicates that regardless of whether or not they are
active, these animals appear to feed only at night (Hammond 1982b), and
food processing is very rapid – particles ingested by the cucumber are
voided from the gut within only about 1 hour!Even when animals are seen apparently feeding during the day,
when collected the researchers found that these animals inevitably had
digestive systems that were completely empty (Hammond 1982b).This is where I come back to the earlier statement that although
these animals are normally nocturnal, they often lose their avoidance of
the light and are sometimes seen cruising around the tank in full
daylight.Although similar studies have not been done on any animals kept
in a reef aquarium for any length of time, studies on natural reefs
indicate that regardless of what looks like normal feeding behavior
during the day, they were not actually ingesting any particles, and were
therefore not feeding,.We do not know why these animals lose their strict nocturnal
schedule in captivity, but there are a couple of likely possibilities.First, the animals may be removed from the predation pressure
that they face on the coral reef, and begin to become bolder over time
in captivity. This is possible, because I have also observed that the
same tends to occur in shallow lagoons and other protected habitats
where, in the absence of predators, these cucumbers are found in high
density during daylight hours.The other possibility is that the animals are starving to the
point that they abandon any semblance of their natural behavior and
start to feed during the daylight.If we assume that it is the former explanation, then (like in
natural populations) the animals may appear to be active during daylight
hours, but are probably not actually feeding.If we assume that it is the latter explanation, then the animals
are doomed to slowly waste away in the absence of sufficient or
appropriate food in the aquarium.
This
is an important distinction, because even if you are fortunate
enough to get a generalist feeder, unless you have a well
established tank with plenty of fine organic detritus upon which
the animal can feed, it is likely to starve to death.These animals are certainly poorly suited to non-reef
aquaria or even traditional Berlin-style reef tanks that are
maintained with a bare or only slightly covered bottom, and from
which detritus is regularly removed.Like most marine invertebrates, these animals are capable
of going for long periods of time without food.Many marine invertebrates can withstand several months or
more of starvation, during which time they slowly shrink while
digesting their internal organs.Depending on the species in question, its initial
condition when brought into captivity, and how often it manages
to locate a suitable food source within the aquarium, many
marine invertebrate species could take more than a year before
they succumb to starvation.
Close-up
of the pinnate feeding tentacles of Synaptula lamperti.
Like most of the apodid cucumbers, one of the
attractions of these animals for aquarists is that these
feather-like feeding tentacles are in almost constant
motion. (Photo by Robert Fenner)
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In
addition to the issue of proper feeding, another important
consideration for getting one of these cucumbers is that these
animals have a highly reduced skeletal system.Like all echinoderms, the skeletal system is composed of a
series of tiny calcareous plates (ossicles) embedded in the skin
of the animal.In
synaptid cukes, these ossicles are reduced to simple hooks called
"anchor ossicles" which project through the skin and
give the animal adhesion to the substrate (and anything else they
touch).Anyone who
has handled one of these cucumbers can vouch for how
"sticky" they are (due to their tiny anchor hooks
snagging into anything they touch), and if you place the animals
into a small container (like an aquarium shipping bag) they often
get snagged even on themselves!Because they lack the tube feet that other echinoderms use
to crawl about, synaptids must crawl about somewhat like an
earthworm, and they use a combination of their muscular
hydrostatic body and the anchor ossicles to crawl around.The muscles in the body squeeze fluids around in much the
same way that a water balloon is deformed when we squeeze it in
our hand.However,
unless there is something to push against, the animal would simply
squeeze out and retract into the same location.The anchor hooks of these cucumbers function in much the
same way as the bristles (chaetae) of a worm they provide traction
against which the animal can push to propel itself forwards.By sticking at their base and “squeezing” their water
balloon body into an elongate shape, they are propelled forward.Then, by hooking in at the front end, and releasing the
grip at the back, when they relax back into the short-n-rounded
water balloon the front end is anchored while the back end is free
to be sucked forward to join it.In this way, their flabby soft body allows them to crawl
about and extend or retract their body with great flexibility.Again, I would refer interested readers to a good
invertebrate zoology textbook, such as Brusca and Brucsa (1990) or
Ruppert and Barnes (1994) for further details.
Their
"half-filled baggie" look is deceptive, however, because
the animals are the fastest and most active of the sea cucumbers,
and are capable of rapidly crawling around the aquarium, or
quickly withdrawing into a crevice when disturbed.The standard feeding behavior of reef-dwelling synaptids is
to anchor about 1/3 of the body into some secure hole, and then
extend the anterior 2/3 of the body to feed (they can do the water
balloon trick with only part of their body if they want to).If something disturbs the animal, it can rapidly contract a
set of muscles that run the length of the body to pull itself back
into that hidey-hole and avoid being eaten.The problem, however, is that those anchor ossicles may be
laying across some of your other invertebrates (such as a coral,
for example), and when the cucumber retracts, it simply tugs those
ossicles free of whatever it was previously anchored upon -- this
can dislodge and/or cause damage to the soft tissue of whatever
the cuke was lying on at the time.Obviously this is a potential concern to aquarists who want
their corals and rock-work to remain in place and undamaged...
So, with those
warnings and caveats, I’ll come back to what I said at the
beginning - these really are one of my favorite odd-ball
invertebrates in a reef aquarium.It’s not because of any service or function they
provide (although some species make excellent grazers or
detritivores), but rather because I think that they make such
an interesting addition (or at least conversation piece) for
my reef aquaria.When
possible, I try to find one of the generalist detritivore
species, such as Euapta lappa, or a diatom grazer such
as Synaptula hydriformis for my tank, because they tend
to thrive in a well-established reef aquarium.If you have a moderately large reef tank with a deep sand bed
and are feeding plankton on a regular basis, there ought to be
plenty of organic detritus for a medusa worm such as one of
these species to find. My cuke spends most of it's time
cleaning the underside of the rocks, and only comes out onto
the surface of the sand and the rocks at night. You can
supplement it's feeding by adding a couple of sinking shrimp
pellets to the area it tends to hang out just before the
lights go out.That
will give the pellets time to soften and fall apart before the
cuke starts to feed, and mine seems to really like the pellet
mush (but so do the hermit crabs, brittle stars, conch, Nassarius
snails and polychaete worms, and they all learn what the
pellets are pretty quickly as well).
The
striking dark-striped white body of Synaptula lamperti
makes an attractive picture against the colored
background of the sponges on which they are
exclusively found. (Photo by Robert Fenner)
Although
attractive to look at, you should resist any urge to
purchase a species of Synaptula because most feed by
mopping tiny particles of organic detritus and
secretions from the surface of living sponges.
Without the correct sponge species on which to live,
they are doomed to a slow death by starvation in your
aquarium.(Photo by Robert Fenner)
If provided with
suitable conditions and sufficient food, some of these animals
may even be able to reproduce in the aquarium.Unlike most echinoderms, sea cucumbers (holothurians)
have only a single, well-developed gonad.In general, the majority of holothurians have separate
sexes, but studies on a couple of synaptid cucumbers suggests
that at least some of them are simultaneous hermaphrodites
(containing both fully functional male and female reproductive
tracts).In the
case of Synaptula hydriformis, researchers found that
these animals are not only simultaneous hermaphrodites, but
they are also capable of self-fertilization (Frick 1998).Even more unusual, the fertilized eggs of S. hydriformis are
retained within the body (in the perivisceral coelomic cavity)
of the adult and actually gain nutrition from the parent,
while they develop internally (Frick 1998).The young are protected and nourished in this way until
they are released as fully-functional juveniles at about 8mm
in length.When
combined with the generalist diatom grazing of this species
(as mentioned earlier), the internal brooding makes for an
ideal candidate for aquarium culture.Other species of synaptid cucumbers may also reproduce
asexually (e.g., Leptosynapta tenuis - Hendler et al.
1995), and again, detritivores capable of asexual reproduction
may make ideal candidates for the aquarium.Individuals of most species also have excellent
regenerative capabilities, so that even when they are injured
accidentally, if they are otherwise healthy and well-cared for
in an aquarium, they are likely to make a full recovery.In fact, Smith (1971a,b) found that healthy animals were
capable of complete regeneration from either the anterior or
posterior end (head or tail) given sufficient nutrient
reserves and appropriate conditions after injury.
Obviously chances of aquarium
reproduction are greatest in species that reproduce asexually
or via brooded offspring.Although roughly 30 species of sea cucumbers brood
their young, most of these are cold-water species, and the
majority of other species spawn their gametes (eggs and sperm)
into the water column to produce larvae.In one study which included four species of
free-spawning synaptid sea cucumbers (Synapta maculata,
Patinapta taiwaniensis, Polycheira rufescens and Opheodesoma
grisea), researchers found that all of these animals tend
to releasetheir gametes late in the summer (Chao et
al. 1995).This
same study found that spawning is correlated with summer
phytoplankton growth, and the authors suggested that direct or
indirect feeding on phytoplankton-derived detritus is a
critical component of nutrition required to reach reproductive
status in these animals.The mode of larval development in most species of
synaptid sea cucumbers remains unstudied.However, for species such as Leptosynapta inhaerens
for which the larval development is known, larvae spend a
considerable time feeding in the plankton prior to
metamorphosing into a tiny version of the adult.For species with this mode of development, successful
reproduction in the aquarium is highly unlikely, and even if a
concerted effort is made to raise the young, the chances of
success are slight (see Toonen 2002 for a detailed explanation
of why this is so, and my Home
Breeders FAQ for details on how this is done).
All-in-all,
however, despite how fascinating these animals are, unless you
are confident of the identification of the animal, can provide
suitable conditions for the animal to feed, and are willing to
live with the potential drawbacks of keeping one of these
animals in an aquarium, they are not really recommended for
keeping in any aquarium.If you have a well-established reef tank, and you take
adequate precautions to protect any pump intakes and/or
overflow drains, and can locate one of the generalist
detritivores that are suitable for the aquarium, I think that
these animals make a fantastic addition to a reef tank,
because they are both active and fascinating to watch.
Although
most of the synaptid cucumbers occur in relatively low
density around coral reefs, Synaptula species can be
quite dense on some sponges. This makes them
easy to collect, but they are not suitable for the
reef aquarium. (Photo by Robert Fenner)
References:
Brusca,
R. C., and G. J. Brucsa. 1990. Invertebrates. Sinauer
Associates, Inc, Sunderland, Mass.
Chao,
S.-M., C.-P. Chen, and P. S. Alexander. 1995. Reproductive cycles of
tropical sea cucumbers (Echinodermata: Holothurioidea) in southern
Taiwan. Marine Biology 122:289-295.
Cunningham,
P., and P. Goetz. 1996. Venomous & Toxic Marine Life of the
World. Pisces Books, Houston, TX.
Delbeek,
J.C., and J. Sprung. 1994. The Reef Aquarium, Vol. 1. Ricordea
Publishing, Coconut Grove, FL.
Frey,
D. G. 1951. The use of sea cucumbers in poisoning fishes.
Copeia:175-176.
Frick,
J. E. 1998. Evidence of matrotrophy in the viviparous holothuroid
echinoderm Synaptula hydriformis. Invertebrate Biology
117:169-179.
Hammond,
L. S. 1982a. Analysis of grain-size selection by deposit-feeding
holothurians and echinoids (Echinodermata) from a shallow reef lagoon,
Discovery Bay, Jamaica. Marine Ecology Progress Series 8:25-36.
Hammond,
L. S. 1982b. Patterns of feeding and activity in deposit-feeding
holothurians and echinoids (Echinodermata) from a shallow back-reef
lagoon, Discovery Bay, Jamaica. Bulletin of Marine Science 32:549-571.
Hammond,
L. S., and C. R. Wilkinson. 1989. Exploitation of sponge exudates by
coral reef holothuroids. Journal of Experimental Marine Biology and
Ecology 94:1-10.
Hendler,
G., J. E. Miller, D. L. Pawson, and M. K. Porter. 1995. Sea Stars,
Sea Urchins, and Allies: Echinoderms of Florida and the Caribbean.
Smithsonian Institution Press, Washington DC.
Kuznetsova,
T. A., N. I. Kalinovskaya, A. I. Kalinovskii, and G. B. Elyakov. 1989.
Structure of synaptogenin B, the artifact aglycone of the glycosides
of the sea cucumber Synapta maculata. Khimiya Prirodnykh
Soedinenii 5:667-670.
Martínez,
M. A. 1989. Holothuroideos (Echinodermata, Holothuroidea) de la region
nororiental de Venezuela y algunas dependencias federales. Boletin
Insitutto Oceanografico Universidad de Oriente Cumana 28:105-112.
Pawson,
D. L. 1986. Phylum Echinodermata. Pp. 522-541 in W. Sterrer, ed. Marine
Fauna and Flora of Bermuda: A Systematic Guide to the Identification
of Marine Organisms. John Wiley & Sons, New York, NY.
Ponomarenko,
L. P., A. I. Kalinovsky, O. P. Moiseenko, and V. A. Stonik. 2001. Free
sterols from the holothurians Synapta maculata, Cladolabes
bifurcatus and Cucumaria sp. Comparative Biochemistry and
Physiology Part B: Biochemistry & Molecular Biology 128B:53-62.
Ruppert,
E. E., and R. D. Barnes. 1994. Invertebrate Zoology. Saunders
College Publishing, Harcourt Brace Jovanovich Publishers, Orlando, FL.
Ruppert,
E. E., and R. Fox. 1988. Seashore Animals of the Southeast: A Guide
to Common Shallow-water Invertebrates of the Southeastern Atlantic
Coast. University of South Carolina Press, Columbia, SC.
Smith,
G. N., Jr. 1971a. Regeneration in the sea cucumber Leptosynapta.I.The process of
regeneration. Journal of Experimental Zoology 177:319-330.
Smith,
G. N., Jr. 1971b. Regeneration in the sea cucumber Leptosynapta.II.The
regenerative capacity. Journal of Experimental Zoology 177:331-342.
Sprung,
J. 2001. Invertebrates: A Quick Reference Guide. Sea
Challengers, Danville, CA.
Toonen,
R. 2002. Aquarium Science: The captive breeding of tropical reef
species for the aquarium trade, with specific attention to long-term
planktotrophic larvae. Tropical Fish Hobbyist #557:66-72.