Reef keeping
aquarists develop an aesthetic judgment about what's “cool” to keep
in their aquariums, and they are often influenced by the rarity of a
particular type of creature. While I too admire a rare specimen, I have
always been and will always be impressed by such common things as
“button polyps,” the many-hued and mostly simple to care for
zoanthids.
Zoanthids form
clusters of polyps and encrusting mats commonly referred to as "sea
mat,""false
coral," or "colonial anemones," and they are sometimes
erroneously called "soft corals." The polyps can be solitary,
connected by a creeping tissue called coenenchyme that may form stolons
or lamellae, or they may be embedded in a thick massive coenenchyme.
They are normally attached to any hard substrate, though some genera are
associated with specific "host" organisms and grow only on
them.
Zoanthids
are relatives of corals and anemones, and are included with them
in the phylum Cnidaria (formerly Coelenterata), in the class
Anthozoa, but the zoanthids comprise the distinct order
Zoanthiniaria (formerly Zoanthidea) in the subclass Zoantharia
that also includes sea anemones, corallimorphs, and scleractinian
corals. Zoanthids differ from true sea anemones, which belong to
the subclass Actiniaria, based on details of their internal
anatomy and the fact that zoanthids form true colonies in which
the individual polyps are connected by a common tissue, the
coenenchyme. True anemones may divide to form beds of adjacent
clones, but they are not connected. Zoanthids lack a fossil
record, but they are considered more closely related to the
extinct "corals" known as Tabulata and Rugosa than to
other anthozoans.
The
order Zoanthiniaria is subdivided into the suborder Brachycnemina,
which includes the families Zoanthidae and Neozoanthidae, and the
suborder Macrocnemina, which includes the family Epizoanthidae and
Parazoanthidae. The structural difference between these groups is
shown in Delbeek and Sprung (1997).
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Genera:
Zoanthids
commonly kept in aquaria belong mainly to six genera, Zoanthus, Protopalythoa, Palythoa,
Isaurus, Acrozoanthus and
a possible new genus for the zoanthid commonly called “yellow
polyps.” There are other genera that are occasionally harvested for
aquaria, including Neozoanthus,
Parazoanthus, and Epizoanthus.
I believe there are also some additional undescribed genera that
occasionally are harvested from Indonesia. One is an extremely small-polyped
Zoanthus-like variety; the
other is a relative of yellow polyps and Acrozoanthus,
with an appearance and colony form somewhere between the two. A third is
quite like this latter one but forms dense colonies like Zoanthus, but with distinctly elongate tentacles. I show this one,
which I call Zoanthus sp? in this article. The solitary polyp zoanthid, Sphenopus,
a solitary polyp species, is not harvested for aquaria, nor is Gerardia,
the only zoanthid known to form a skeleton. The genera Thoracactis and Isozoanthus
in the family Parazoanthidae are also poorly known, and not likely to be
seen in an aquarium.
Acrozoanthus
aka ³Stick Polyps² grow on the parchment-like tube of a
polychaete worm.
The author found this unidentified, large, non-photosynthetic
zoanthid in the Solomon Islands during a night dive. It was under
a ledge and the tentacles were expanded.
Most
sea anemones sting and injure zoanthids.
Typical
daytime appearance of "Snake Polyps," Isaurus
tuberculata. Sometimes the columns are fluorescent green or
have iridescent metallic spots.
The
sundial snail Heliacus feeds on zoanthids.
This
recently imported Zoanthus sp. is unusual in that it has a
very thick coenenchyme in which the polyps are embedded.
Open
polyps of Neozoanthus.
Closed
appearance of the uncommon zoanthid Neozoanthus growing on
the base of a Sarcophyton sp. leather coral. Note sand
grains attached to columns.
The
Hawaiian Protopalythoa psammophila is a wonderful shade of
green. Similar forms occur in the Sea of Cortez and Singapore, and
other species may have this same coloration.
Protopalythoa
vestitus has a distinctive
pattern of stripes. Protopalythoa grandis has a rare form
with similar stripes.
Palythoa
caesia
from the Solomon Islands.
This
Palythoa sp. has fluorescent orange pigment, unusual for
the genus.
Parazoanthus
axinellae
from the Mediterranean. A beautiful temperate species that grows
freely on rocks, unlike most other members of the genus that
associate with sponges and other animals that provide a position
for them out in strong currents where they can catch plankton.
Reproduction:
Ryland, and
Babcock (1991) describe the reproductive cycle of Protopalythoa
on the Great Barrier Reef in Australia. There it spawns along with
corals during the mass spawning event during the week after the full
moon in November. Zoanthids may have separate sexes, but some are also
hermaphrodites (Fosså and Nilsen, 1998). The product of the union of
gametes develops into a free- swimming larva called Zoanthina for Zoanthus
spp., and Zoanthella for Protopalythoa
spp (Delbeek and Sprung, 1997).
Competitive
interactions:
Zoanthids
compete for space as do most other sessile encrusting or colonial
invertebrates, and they do so by means of toxins or the ability to
rapidly overgrow their neighbors. Palythoa
and Protopalythoa are toxic
and will damage most stony corals and some soft corals. Zoanthus species are more benign, but often overgrow their
neighbors. Zoanthus spp. are
easily stung by the sweeper tentacles of many scleractinian corals, and
also by sea anemones. Zoanthus
spp. generally are safe to place adjacent to most soft corals, however.
Yellow polyps should not be placed where they will contact soft corals
of the genus Clavularia. Yellow polyps, Zoanthus,
Protopalythoa and Palythoa spp. can all be grown adjacent to each other with no harm
done to any of them.
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Zoanthus:
Members of
this genus are the most colorful of the zoanthids, being shades of
green and brown typically, but sometimes fluorescent red, orange,
pink, lavender, blue, yellow, or gray, and usually two-toned. They
form colonies of densely crowded polyps attached in a common
tissue at the base. Some varieties form branched stolon-like
creeping bases that tightly adhere to the substrate surface. One
form recently imported for aquaria has polyps imbedded in a thick
coenenchyme, much like Palythoa spp. Many species grow in the intertidal zone, being
exposed at low tide to the baking sun, cold fronts, or to rain
showers. They slowly release the water they retain in their
columns in order to prevent either desiccation or damage from
freshwater during a heavy rainstorm.
Zoanthus
spp. do not need to be fed directly, since they obtain much of
their nutritional requirements from their symbiotic zooxanthellae.
They must therefore be provided with adequate illumination to
thrive. They also ingest dissolved organic substances from the
water, as well as fine particulate matter. Some species do not
take large particles of food, while others do take and eat such
things as flake food, blackworms, shrimp, and sea urchin eggs.
Regular feeding promotes rapid growth.
In
the Caribbean there are two species commonly known and described,
Zoanthus sociatus and Z.
pulchellus. In south Florida there are two additional species
apparently never formally described. One of these undescribed species (Zoanthus
n.sp. 1 in my photos) lives only intertidally, on rocky jetties at
inlets where the ocean feeds into the intercoastal waterway. It forms
continuous thick sheets of coenenchyme with the polyps imbedded, and the
colors of the polyps sometimes include fantastic combinations in bright
hues. This species is commonly mixed with Z.
sociatus and Z. pulchellus
where it occurs, and it is easily distinguished from them. The other
undescribed species (Zoanthus
n.sp. 2 in my photos) forms very small colonies of just a few to perhaps
a few dozen polyps, usually on hardbottom reefs in sandy areas. The
polyps are spaced widely and connected by a stolon. They are typically
bright green, but may also be dark blue-gray or sometimes brilliant
turquoise blue. The polyps are small and flattened, considerably smaller
in diameter and height compared to Z.
sociatus. Zoanthus sociatus
forms large colonies of polyps connected at their bases by a very thin
coenenchyme, so loosely connected are they that individual polyps or
clumps of polyps readily break off of a colony if tugged. The coloration
is variable but is generally shades of green or blue-green, often with a
yellowish oral cone. Zoanthus
pulchellus is found mainly on offshore coral reefs, though it can
occur in inlets. It has larger polyps than Z.
sociatus, and perhaps larger than any other species of Zoanthus.
Zoanthus pulchellus readily
accepts food while Z. sociatus does
not. Zoanthus pulchellus is
also more variable in coloration, being shades of green, gray, pink, or
fluorescent red. Colonies of this species may cover large areas of reef
substrate, particularly on back reef rubble.
Parazoanthus
sp. growing on a sponge.
Parazoanthus
spp
Parazoanthus
spp
2
A
single polyp of Protopalythoa. Note the tapering tentacles. The
tentacles in Zoanthus are by comparison usually shorter and blunt
tipped.
A
fluorescent red Zoanthus sp. on a reef slope in the Solomon
Islands.
Blue
zoanthids are highly prized by zoanthid lovers. This little patch
of gems was among other colors growing on a reef crest in the
Solomon Islands.
Zoanthus
sociatus from Florida. It
requires bright light.
Yellow
polyps have not yet been classified, though they have been popular
in aquariums for nearly 30 years!
This
undescribed Zoanthus sp. from South Florida forms sheets
covering rocks in places with strong currents, such as inlets.
This one is a nice combination of pink, blue, and green
This
undescribed Zoanthus sp. from South Florida grows in small
colonies with the polyps connected by a stolon-like coenenchyme.
Several species of
Zoanthus are commonly imported
for the aquarium trade mainly from Indonesia and Singapore. Based on
their similarity to two of the Caribbean species it seems to me that
some of them may have circumtropical distributions, though I have not
seen anything in the literature to suggest that is the case.
Protopalythoa:
Members of this
genus were previously referred to as Palythoa
spp. in scientific and aquarium literature. Muirhead and Ryland (1985)
separates this genus from Palythoa,
which has its polyps deeply embedded in the coenenchyme. The fact that
there are species with intermediate structure, such as Palythoa
mammillosa, suggests that this distinction is of questionable
significance. Borneman (1997) also discusses this issue. Nevertheless
the majority of species of Palythoa
can readily be distinguished from Protopalythoa
based on this characteristic. Protopalythoa
spp. have the largest polyps of all the zoanthids. They have slimy thick
tissue, often with some sand grains trapped on the lower portions of the
columns of each polyp. Protopalythoa
spp. may occur on reef slopes, but are most common on sandy reef flats
and hardbottoms in shallow or deep water. Most species are brownish,
sometimes with green oral discs. Some varieties have striped or mottled
oral discs. These zoanthids always remind me of the insect-eating plant
known as “Venus flytrap,” since they have tapering tentacles around
the margins of the oral disc quite like the pointed “eyelashes” that
fringe the food capturing structures of the aforementioned plant. The
plant and zoanthid polyps also slowly close around their prey in a
similar fashion. Protopalythoa
spp. have symbiotic zooxanthellae and must be kept under moderate to
strong illumination to thrive.
Protopalythoa
spp. have toxic mucus that is able to destroy scleractinian coral
tissue, so be careful not to place colonies where they are likely to
contact stony corals. I'll discuss this toxic mucus in more detail later
in the article.
Protopalythoa
spp. all take food when it is offered. They feed on meaty things such as
shrimp, worm, or mollusk flesh, and will also take fish eggs,
invertebrate eggs, flake foods, and pellet foods.
Three easily
identified species are worth distinguishing from the rest, Protopalythoa
grandis, P. psammophila,
and P. vestitus. My favorite
is Protopalythoa grandis,
which has polyps up to two inches in diameter, and mottled in shades of
brown, white, and green. It apparently has circumtropical distribution
(Colin and Arneson, 1995), but colonies harvested for the aquarium trade
come mainly from deep hardbottom reefs in the Gulf of Mexico off the
Florida coast. To grow this species it is necessary to offer food
regularly. Without adequate food supply the polyps shrink slightly and
the fringe of tentacles degenerates to the point that the polyps could
be confused with Discosoma spp.
Protopalythoagrandis
has a habit of closing up for several days, during which time it
secretes a waxy film that is subsequently shed. This is a normal process
that is also seen in Palythoa spp., other (but not all!) Protopalythoa spp., as well as in leather corals such as Sarcophyton
and Lobophytum, gorgonians such as Pterogorgia,
and the stony coral Porites.
Shedding this waxy film is a means of preventing the growth of algae or
other organisms on the surfaces of the polyps. Protopalythoagrandis should be placed near
the bottom of the aquarium where the illumination is not very intense.
Protopalythoa
psammophila ranges
across the Pacific and has the desirable feature of being bright
fluorescent green. At one time this species was harvested from Hawaii
for the aquarium trade, but such harvest ended in the early 1990's due
to new regulations for the state. Many aquarists still grow and
propagate the Hawaiian form, but bright green Protopalythoa
cf. psammophila are also now
occasionally being imported from Singapore and the Sea of Cortez. This
species prefers bright illumination, but the most intense color develops
under moderate illumination.
Protopalythoa
vestitus is a beauty
that is widespread in the Indopacific. It has a distinctive pattern of
brown and white bands on the oral disc, and may fluoresce green under
blue light. It is commonly imported from Indonesia.
Palythoa:
Members of this
genus all look more or less the same, quite like faviid corals with
which they are often confused. As mentioned previously, the coenenchyme
is thickened into a cushion in which the polyps are embedded. The
coenenchyme also contains sand grains. The color is usually pale yellow
or brown, but fluorescent green colonies occur in some regions, and I
observed some with fluorescent orange pigment in the Solomon Islands. A
small colorful crab, Platypodiella,
is associated with this zoanthid and apparently feeds on it, potentially
obtaining a benefit from the toxin(s) the zoanthid produces (Delbeek and
Sprung, 1997).
Palythoa
spp are most common on reef flats or the tops of reefs, where they may
form colonies several metres across. They often occur in a zone mixed
with fire corals (Millepora
spp.). They may also occur on reef slopes, usually as small colonies.
Degraded reefs may become overgrown by them, and they are able to
out-compete stony corals.
In the aquarium
one should be careful not to place Palythoa
where it will contact stony corals. Palythoa
spp. grow best under strong illumination, and should be offered
planktonic foods, such as copepods, Artemia,
or Daphnia.
Isaurus:
Members of this
genus are commonly called "Snake Polyps.” Three valid species are
known, I. tuberculatus, I.
cliftoni, and I. maculatus.
Isaurus tuberculatus is distributed circumtropically and is the
species most commonly seen in the wild and in aquariums. Isaurus spp. have elongate columns with tubercles (bumps) and the
columns are typically oriented so that the ends face downward. The
tentacles are normally retracted during the day and expand at night to
feed on zooplankton. The columns contain zooxanthellae and should be
exposed to bright light. Isaurus
does not need to be fed to be maintained in captivity, but regular
feeding will cause the polyps to multiply. The best foods for Isaurus
are Daphnia, Cyclops (copepods), and brine shrimp nauplii.
Yellow
Polyps:
Although various
aquarium references have called this zoanthid a Parazoanthus
sp., it is not. That description makes sense when one considers that Parazoanthus spp. are often yellow. For example, several years ago
when I discussed "the yellow photosynthetic zoanthid from
Indonesia" with world zoanthid expert J. S. Ryland, he assured me
that it was probably a Parazoanthus,
but he cautioned that his comment was based on the color I described
only, not on a specimen. This zoanthid does not yet have a name, nor
even a genus. It appears to be related to Acrozoanthus
and the undescribed zoanthid I am calling Zoanthus
nsp.? in this article.
Despite its lack
of taxonomic placement, this zoanthid has had a place in aquariums since
the very beginning of the reefkeeping hobby. It is a very hardy
beautiful addition to any aquarium. The only downside is that it spreads
quickly enough to damage and overgrow corals. Take care to place this
species on isolated rocks in the sand or on the glass to avoid having it
grow on the reef and approach corals. Care of yellow polyps is simple:
give them strong light and feed them. If they are placed in strong water
motion they will catch the food offered to the fishes, and do not need
supplemental feeding. They eat flakefood, shrimps, worms, or anything
meaty and small enough for them to grab.
Acrozoanthus:
One species in
this genus is frequently imported under the common name “Stick
Polyps.” The name arises from the parchment like “stick” on which
these polyps occur naturally. The stick is not secreted by the polyps
themselves, but is instead the top of a tube formed by a large type of
polychaete worm (Eunice tubifex)
that lives buried in sand. The collectors simply cut off the top of the
tube, leaving the worm intact in its home buried in the sand.It can readily build a new extension of its tube upon which more Acrozoanthus
can settle.
Acrozoanthus
polyps are similar to those of “Yellow Polyps,” but they are larger,
brown, and have very long stringy tentacles. Though naturally associated
with the worm, these polyps readily adapt to captivity and spread onto
rocks. They need to be fed to really proliferate in captivity. They will
eat copepods and any chopped meaty foods. Despite their need for food,
they do also harbor zooxanthellae, so they should be maintained in a
well illuminated aquarium.
Neozoanthus:
This
uncommonly seen genus forms small colonies of cryptic polyps that are
similar in appearance to those of Epizoanthus.
They characteristically have sand grains trapped in their tissue. They
should be fed with copepods or brine shrimp nauplii, and provided with
moderate to strong illumination. I first observed this species as a few
polyps on the base of a Sarcophyton
sp. from Indonesia.
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Parazoanthus
and Epizoanthus:
Members of these genera are
epizootic, meaning they live on other animals. Parazoanthus
and Epizoanthus are almost
exclusively found associated with sponges and hydroids, though they
may also grow on gorgonian skeletons and even freely. They do not
harbor zooxanthellae and thus must be fed zooplankton regularly in
order to thrive. They occur where there are strong currents that carry
plankton, so the water motion in the aquarium housing them should be
swift and laminar. The association with sponges is believed to protect
the sponges from predators. These zoanthid polyps are toxic and may
also sting sponge eaters such as large angelfish.
Predators:
Some fishes will eat
zoanthids, though most find them distasteful. The box snails, Heliacus
spp., feed specifically on zoanthids, and can be considered a pest if
one is trying to grow them, or a savior if one is trying to rid the
aquarium of zoanthids. It is not always easily seen, as the shell is
round and about the size of a closed zoanthid polyp.
A
word of caution about Zoanthids:
I must mention the fact that
zoanthids are quite toxic. They produce a substance known as palytoxin
(Mebs, 1989) that is one of the most toxic naturally occurring poisons
known. This substance was first discovered associated with Palythoa
spp. in Hawaii, but has since been found in Zoanthus
as well (Fosså and Nilsen, 1998). Palytoxin is apparently produced by
bacteria that live in association with zoanthids. A fascinating
account about the discovery of palytoxin can be found online on the
Wet Web Media site, see reference given.
A strange side note to this
is the anecdotal observation I reported in my column Reef
Notes in FAMA magazine that this toxin or another one associated
with zoanthids may be able to be aerosolized. An aquarist trying to
rid his live rocks of a species of Protopalythoa
decided to remove the rocks and spray boiling water on them to kill
them. A friend of his contacted me after the aquarist was in the
hospital and in serious condition, the doctors unable to determine
what had caused a serious reaction and respiratory distress. I pointed
out the possibility of a palytoxin reaction, but was skeptical about
the aerosol or "toxic fumes" that the aquarist believed made
him become ill suddenly. The aquarist later recovered, but slowly. In
any case, one should be extremely careful when handling zoanthids, Protopalythoa
and Palythoa spp., in
particular. Rinse your hands thoroughly with soap and water
immediately after contacting them.
Burnett,
W.J., et. al. 1997. Zoanthids (Anthozoa, Hexacorallia) from the Great
Barrier Reef and Torres Strait, Australia: systematics, evolution and a
key to the species. Coral Reefs
16: 55-68.
Coll,
J.C., and P.W. Sammarco. 1988. The role of secondary metabolites in the
chemical ecology of marine invertebrates: a meeting ground for
biologists and chemists. Proc. 6th Int'l Coral Reef Symp, Australia, 1: 167-73.
Delbeek,
J.C. and Julian Sprung. 1997. The
Reef Aquarium. Volume 2. Ricordea Publishing,., 546 pp.
Fosså, S. and A.
J. Nilsen (1998). The Modern Coral Reef Aquarium Volume 2. Birgit Schmettkamp Verlag
Koehl,
M.A.R. 1977. Water flow and the morphology of zoanthid colonies. Proc.
3rd Int'l Coral Reef Symp. 437-44.
Mather, P. and I.
Bennet (eds.) (1993). A Coral Reef Handbook 3rd Ed. Surrey Beatty and Sons PT
Ltd., Chipping Norton, NSW Australia.
Mebs, D. (1989). Gifte
im Riff. Wissenschaftliche Verlagsgesellschaft mbH, Stuttgart,
Germany.
Muirhead and
Ryland (1985). A review of the genus Isaurus Gray, 1828 (Zoanthidea),
including new records from Fiji. Jour. Nat. Hist. (19): 323-335.
Ryland, J.S. and
R.C. Babcock (1991). Annual cycle of gametogenesis and spawning in a
tropical zoanthid, Protopalythoa sp. Hydrobiologia
(216/217): 117-123.
Sebens,
Kenneth P. 1977. Autotrophic and heterotrophic nutrition of coral reef
zoanthids. Proc. 3rd Int'l Coral
Reef Symp. 397-404.
Sprung, J. (2000) Invertebrates:
A Quick Reference Guide. Ricordea Publishing. Coconut Grove, Fl.
USA.