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EDITORS PAGE by James M. Lawrence



RARITIES Pseudocoris Wrasses by Scott W. Michael

by Daniel Knop
by Daniel Knop

by Claude Schuhmacher

by Daniel Knop

by Nate Wilson

by Ronald L. Shimek, Ph.D.


by Erik Stenn

by Willi Wehner
Japanese Swallow Angelsh by Daniel Knop

Heavy-metal poisoning by Daniel Knop;
Flame Hawksh by Inken Krause
CORAL SOURCES: Outstanding aquarium shops
CORALEXICON: Technical terms that appear in this issue
World-class aquarium shops & places to visit

High drama for the Hawaiian aquarium trade by Ret Talbot
REEF LIFE: by Denise Nielsen Tackett and Larry P. Tackett
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, The Reef & Marine Aquarium Magazine

(ISSN:1556-5769), is published bimonthly in January,
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is a licensed edition of KORALLE Germany,

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Zoanthids, Zoanthus sp. Photo: D. Knop.
Background: Blue Zoanthus sociatus
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notes f rom DANI EL KNOP
still remember well a lecture that reef aquarium
pioneer Peter Wilkens gave 25 years ago. The place:
Mannheim; the subject: Encrusting anemones in
the marine aquarium. He explained in minute de-
tail what one needed to know about their aquarium
maintenance, and his large medium-format slides
showed the individual color morphs that were avail-
able back then. There werent many of them, and most
were shades of brown and olive green. But these ower-
like actinians, with their color-contrasting mouth discs
and tentacles, were fascinating, undemanding, and ro-
bust, so they often thrived in captivityeven in those
early years of the reef aquarium hobby, when conditions
were anything but optimal in many respects. So theyve
been around right from the start.
Encrusting anemones remained an ideal and popu-
lar livestock for beginners throughout the entire 1980s,
but then suffered an abrupt descent into a wallower
existence; quite simply, people found colorful stony cor-
als more attractive. In addition, many considered the
difculty of stony-coral keeping a challenge; it was new
territory, and that made it alluring. After all, everyone
had zoanthids. A st-sized substrate rock with a thick
cushion of polyps often cost less than a tiny twig of a
colorful Acropora.
But the wind has changed once again. The time of
bargain-basement prices is nearly over. Numerous color
morphs of encrusting anemones, endowed with inven-
tive names, are offered for sale. Polar Ice Caps, Bali
Crackers, and Cherry Mints have garnered fans
worldwide. Keeping their fantastic pigmentation also
represents a certain challenge for both aquarium equip-
ment and aquarist. And they are ideally suited to small
nano reef tanks with medium-strength lighting, which
should cause uorescent effects to appear and at the
same time save on electricity. This should guarantee their
popularity for years to come.
Happy reading!
FOCUS: Hoodwinked
Stony coral and soft coral in peaceful coexistence? Anyone who
thinks the animal on the left is soft coral has been hoodwinked
by the rened mimesis of a nudibranch. This adaptation to the
specic coral species on which the gastropod feeds camouages
it from its predators and makes life difcult for them. Interest-
ingly, however, it was those very predators who brought about
this fantastic mimicry: predator pressure continually sorts out
poorly adapted gastropods.
Scientifcally undescribed reddish-
brown nudibranch of the genus
Marionia, family Tritoniidae. Favia sp.
moon coral, right.
KORALLE editor Daniel Knop
documents the development of
young nudibranchs feeding on
encrusting anemones.
y personal quest for this extraordi-
nary sh began just over 20 years ago,
when I received a photo of an unusual
discovery taken by a diver friend, Kal
Muller. Kal took the photo during his
recent visit to a remote is-
land off the eastern coast
of Sulawesi. The wide-angle shot showed a
group of apogonids sheltering near a Long-
Spined Sea Urchin. It was denitely some-
thing special. In fact, I had never seen such
a spectacular cardinalsh and assumed it
certainly must be new to science. Somehow
I would have to nd a way to visit the Bang-
gai Islands and collect this fantastic sh!
It took two more years before I was able
to arrange travel to the Banggai Islands, as
a side trip in conjunction with a biodiver-
sity conference I planned to attend at Ma-
nado, in northern Sulawesi. Timing would
be tight as there were only two ights per
week to Luwuk, the jumping-off point to
the Banggai Group. I invited frequent div-
ing companion and renowned underwater
photographer Roger Steene to join the mini-
expedition. The trip was arranged for mid-
November 1994. We would y to Luwuk on
the Thursday ight, make a quick visit to
the Banggai Islands, hopefully collect and
photograph the sh, and return to Manado
on the Sunday ight. It didnt leave much margin for
error, but the busy conference schedule didnt allow for
extra time.
We arrived at Luwuk around noon and spent most
of the day arranging passage on the Banggai ferry and
shopping for snack foods. The ferry nally departed at
midnight and we prepared for a sleepless night on deck
with the throng of about 100 passengers, vehicles, and
livestock. But to our pleasant surprise we were able to
bargain with the captain, negotiating the hire of his per-
sonal quarters for 50,000 rupiahs, probably more than
he would earn in salary for the entire voyage. The cabin
was very small, but nevertheless comfortable. There were
two beds, a fan, and an adjoining toilet, and room to
spread out the photographic equipment. The journey
took 12 hours, but the time passed quickly, especially as
we were able to sleep in relative comfort. The last hour
of the voyage was spectacular, as the ship negotiated a
narrow passage between two jungle-clad islands. At last
the vessel docked at the main wharf at Banggai. It was
12:00 noon and time to put our much-discussed plan
Banggai Rescue Proj ect: A bi ol ogi sts perspecti ve

Dr. Gerry Allen with frst live specimens of
Banggai Cardinalfsh he collected in 1994.
For many of us, Dr. Gerald R. Allen, known throughout the aquatics world as Gerry, is one of the living heroes of reef
sh science. A protg of the legendary Dr. John Randall, Gerry has personally found and described a tremendous
array of new sh species and published a wide-ranging library of marine guidebooks and scientic papers, while per-
severing in efforts to preserve marine species diversity in the Coral Triangle. As the ichthyologist who introduced the
Banggai Cardinalsh to modern science and the aquarium hobby, Gerry has graciously lent his support to The Banggai
Rescue Project. Here is an excerpt from his Foreword to the soon-to-be-published Banggai Cardinalsh book, coming
from the publishers of CORAL.
James Lawrence
Shelburne, Vermont
into action. We didnt have
any time to spare; the ferry
would depart in six hours.
Race to the pearl oyster farm
We hired the ferrys radio op-
erator, whom we nicknamed
Sparky, to accompany us
because he spoke a few words
of English. As soon as the ferry
was securely tied Sparky went
ashore to hire a small motor-
boat, and within 30 minutes
we were headed south along
the western side of the island.
Kal had given us vague instruc-
tionshe had found the sh near a wooden jetty at a
pearl oyster farm owned by a Chinese man, in a bay
about one hour by motorboat south of Banggai town.
Sparky relayed this information to our driver, who nod-
ded in recognition at the mention of orang cina, the
Indonesian translation of Chinese man. Several pearl
oyster farms are located along the coast, but evidently
only one is owned by a Chinese person. Forty-ve min-
utes later we pulled in to a narrow wooden jetty at one
side of a picturesque bay.
It was a race to be rst in the water. In less than a min-
ute we were both submerged, but the sh was nowhere to
be seen. I nned slowly away from the jetty, methodically
checking every square metre. Kal had previously located
the sh in only 6.5 feet (2 m) of depth, so the search was
limited to shallow water near the shoreline. The bottom
was an uninteresting blend of sandy silt and clumps of
seagrass. After a dozen breath-hold dives I sorely missed
the luxury of our usual scuba equipment. I inhaled an-
other big breath and plunged down. Swimming close to
the bottom, I rounded a large patch of seagrass, and sud-
denly there it wasa group of about 10 adults huddled
around a long-spined Diadema sea urchin.
Its difcult to describe the level of excitement at
that moment, but sufce it to say there was a maximum
adrenalin surge. The beauty of this sh in its natural
habitat is something to behold. The combination of a
striking color pattern and long, graceful laments on
the dorsal and tail ns is truly spectacular.
We had to work fast. I calculated we should spend
no more than three hours at the site to allow ourselves
ample time for the ferry departure. Further searching
revealed several more groups, invariably huddled close
to urchins, including a large aggregation containing
more than 50 sh. Our rst priority, and the most time-
consuming chore, was underwater photography. Over
the next two hours we took more than 200 shots. This,
of course, was the predigital era, so after each 36-shot
roll we had to tediously exit the water, towel off, and
change lm. Finally, with only half an hour remaining,
it was time to collect a small sample. This
proved a simple task, as the sh retreated
among the sea urchin spines where they
could be easily sandwiched between a
pair of small hand nets. The rst attempt
yielded six adults, which were summari-
ly placed in a plastic bag. One of the sh
spat out an orange-coloured egg mass
not unusual, as male cardinalshes
are well known for their habit of oral
egg incubation. A few more specimens
were captured and placed in a separate
bag. I could scarcely believe my eyes
when I checked the second bag a few
minutes later. There were more than
two dozen miniature replicas of the adult sh that ap-
parently had been expelled from the mouth cavities of
two large sh that appeared to be incubating eggs, judg-
ing from their swollen throats. Brood care of live young
was previously unknown in cardinalshes.
Back aboard the ferry, I carefully pinned out the ns
of the collected sh and preserved them in formalin so-
lution for later study. We were ecstatic that our care-
fully laid plans had unfolded with clockwork perfection.
Not only had the sh been successfully pho-
tographed and collected, but we also gained
a sneak preview of its unusual lifestyle and
breeding habits. To make things even sweeter,
I thought this amazing sh was a new scien-
tic nd. However, detailed examination of
the specimens in my laboratory at the West-
ern Australian Museum and a review of taxo-
nomic literature proved this assumption to be
wrong. The sh had already been described!
My investigations revealed that two subadult
specimens were collected at Banggai Island in
1920 by a Dutch physician named Kaudern.
The specimens had been sent to the Natural
History Museum in Leiden and the species
was eventually described in 1930 as a new ge-
nus and species, Pterapogon kauderni.
Sudden limelight for an obscure species
It seems hard to believe that this magnicent
sh escaped the attention of collectors for de-
cades, considering that Indonesia is a leading
exporter of marine shes for the international
aquarium trade. However, it remained elusive
thanks to the lack of a pelagic dispersal stage
typical of most reef shes and the consequent
extremely limited geographic range, conned
to an area seldom frequented by outsiders.
Suddenly the Banggai Cardinalsh was thrust
into the limelight, becoming an overnight sen-
sation. I recounted the tale of its rediscovery
at the Louisville MACNA Conference in 1995,
it w
two dozen miniat



and again in an article that appeared in the May 1996 is-
sue of Tropical Fish Hobbyist Magazine. This was followed
by a brief scientic paper reporting our observations of
its natural habitat and unusual oral brooding habits.
I have experienced a certain degree of guilt for having
triggered interest in this species, which almost overnight
became one of the most popular species in the aquarium
hobby. Considering its limited distribution, I was partic-
ularly disturbed to discover that thousands of specimens
were being captured and exported each monthnot an
ideal conservation scenario for a sh that is geographi-
cally restricted and produces relatively few eggs com-
pared to most reef shes.
It is therefore particularly gratifying to see the initial
results of the ongoing Banggai Rescue Project presented
in this book. Hopefully, this welcome addition to our
knowledge of this fascinating species will lead to reforms
of the overshing situation in the Banggai Islands and a
solution to the bafing iridovirus problem that has had
such a severe impact on imported specimens in recent
years. Importantly, this book also includes the latest
information for successfully rearing and maintaining
Banggai Cardinalsh in captivity, a positive step that will
certainly reduce the demand for wild-caught sh, thus
making a valuable contribution to the conservation of
the natural population.
Gerald R. Allen, Ph.D.
Perth, Western Australia
Addendum: On a subsequent visit to the islands in 1997
I had an opportunity to dive in Banggai Harbor, includ-
ing next to the ferry jetty. To my surprise the Banggai
Cardinalsh was exceedingly abundant among the jetty
pylons and elsewhere around the harbor. Had we known
this, we could have saved lots of time and energy, not to
mention angst, on our initial 1994 visit.
The Book: Now in its nal stages of production, the
Banggai Cardinalsh book (opposite page) will be pub-
lished in the spring of 2013. Starting with this excerpted
Foreword and continuing with a rst-hand look at the
Banggai Cardinalsh in its native habitat, the book cov-
ers the shs natural history, conservation status in the
wild, reproductive habits, and ways for small-scale breed-
ers to become local suppliers of captive-bred Pterapogon
kauderni. To sign up to receive notice of the publication
date and to order the book, visit: http://www.banggai-
Local fshers moving wild-caught Banggai
Cardinals to a holding pen in Sulawesi,
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The purple and pink sunscreens of stony
corals and how they work
New research reveals that corals use their pink and pur-
ple hues as sunscreen to protect them against harmful
Shallow-water coral reefs thrive in blazing tropical
sunlight, as they benet from sugars and lipids that are
produced by their light-dependent zooxanthellae (symbi-
otic algae). However, in the upper reaches and shallow-
est areas of coral reefs, light levels are often higher than
required by the corals, so paradoxically, the vital sunlight
can become harmful for the algae and their hosts.
Apart from temperature, light stress is a major driver
of coral bleaching, the loss of zooxanthellae that can
lead to loss of coral tissue and death.
Working on the Great Barrier Reef and under tightly
controlled conditions in the Coral Reef Laboratory of the
University of Southampton, a team of researchers pro-
duced experimental evidence that the pink and purple
chromoproteins can act as sunscreens for symbiotic al-
gae by removing parts of the light that might be harmful.
Dr. Jrg Wiedenmann, senior lecturer of biological
oceanography and head of the Universitys Coral Reef
Laboratory, who led the study, says: The beautiful pink
and purple hues that are produced by the coral host are
often evoked by chromoproteinspigments that are
biochemically related to the green uorescent protein
(GFP) of the Crystal Jellysh, Aequorea victoria. In con-
trast to their green, glowing counterparts, the chromo-
proteins take up substantial amounts of light, but they
dont re-emit light.
GFP-like proteins were suggested to contribute to the
f i ndi ngs and happeni ngs of note i n the mari ne worl d
Purple Acropora valida
displaying light-protective
protection of corals and their symbionts from excess sun-
light. This hypothesis has been controversially discussed,
as the mechanism as to how these pigments function re-
mained unclear. At least for the chromoproteins, we know
now that they have the capacity to fulll this function.
The researchers also proposed an explanation for the
mysterious phenomenon that some corals accumulate
exceptionally high amounts of chromoproteins in grow-
ing areas, such as branch tips or in the region of healing
Dr. Wiedenmann, who is based at the National
Oceanography Centre, Southampton, explains: These
growing areas contain essentially no symbiotic algae, so
much of the light is reected by the white coral skel-
eton instead of being used by the algae. The resulting
increased light intensities in the new parts of the coral
represent a potential danger for the algal cells that need
to colonize these areas. Hence, it seems that the corals
use a clever trick to help their symbionts. The higher
light intensity switches on the genes that are responsible
for the production of the sunscreening chromoproteins.
Our results suggest that the screening effect of the
chromoproteins could help the algae to enter the new tis-
sue. Once the symbiont population is fully established,
the light levels in the tissue decrease, as the algae use
most of the light for photosynthesis. As a consequence,
the genes of the chromoproteins are switched off again,
which allows the coral to save the en-
ergy required for their production.
The research contributes to a
better understanding of the corals
response to environmental stress.
Knowledge of the stress resilience of
corals is an important requirement
to help predictions of the fate of cor-
al reefs that are exposed to climate
change and various forms of anthro-
pogenic disturbance.
From materials released by the University
of Southampton and the researchers paper published in the
latest edition of the journal Coral Reefs, January, 2013.
Reef speedster wrasse challenges tuna in
ocean sprints
Australian scientists working on the Great Barrier Reef
have found that modestly sized coral reef wrasses can
swim as fast as some of the swiftest shes in the ocean,
using only half as much energy to do so.
By apping their ns in a gure-eight pattern, Redspot
Ribbon Wrasses (Stethojulis banandensis) can travel at
high speeds while using 40 per cent less energy than Yel-
lown Tuna. (This wrasse species is known in Australia as
the Bluelined Wrasse, and in other ref-
erences as the Red Shoulder Wrasse.)
For a long time, people thought the
best high-speed swimmers were the sh-
es cruising in open waters, like mackerel
and tuna, says Dr. Chris Fulton of the
ARC Centre of Excellence for Coral Reef
Studies and The Australian National Uni-
Our study shows that these coral
reef wrasses, by virtue of their unique
wing-like ns, can maintain very similar
speeds at a dramatically lower energetic
cost, he says.
The researchers discovery could help
revolutionize robot submarine technol-
ogy by reducing the amount of energy
needed to propel objects underwater.
Current Autonomous Underwater
Vehicles (AUVs) use propellers or jets at
the back. By replacing these with ns at
the front to mimic how the ribbon wrass-
es ap their ns, we could propel robots
with less power, saving on batteries and
increasing their range, Dr. Fulton says.
Dr. Fulton explains that shes like
tuna and mackerel move their bodies
and tails to propel themselves through
the water. While this method enables
them to swim fast, it can come at a high
energetic cost.
Another way shes swim is to use
their pectoral ns, those at the front of
their bodies, to produce thrust, he notes.
Fishes that do this with rounded ns tend
to paddle their ns back and forth, almost
the way we row a boat: they hold their ns
out and pull back in a power stroke, then
collapse their ns and bring them forward
for a recovery stroke. This means they are
producing thrust only half the time.
Redspot Ribbon Wrasses, however,
ap their tapered ns in a gure-eight pattern that pro-
duces thrust on every stroke, making it far more energy
This gure-eight n sweep allows these wrasses to
create a lift force as the water ows over their ns, in a
very similar way to how birds y through the air. This
means the sh are literally ying underwater.
They also hold their bodies rigid while swimming
to make them as streamlined as possible. They only ap





Redspot Ribbon Wrasse (Stethojulis banandensis), terminal phase
male. These fsh can outpace athletic tunas using 40 percent less
their ns, slightly adjusting the angle so as to cruise
along without burning up a lot of energy.
The study shows that Redspot Ribbon Wrasses stand
out as the highest-performing swimmers for their size
with respect to optimum swimming speed and energy
consumption, Dr. Fulton says.
Such extreme performance appears to be linked with
the shs habitat, he says. Unlike many tail-swimmers
that dwell in the open ocean, these wrasses live on shal-
low coral reefs, where they experience rough treatment
from waves breaking over the reef.
Most people think that coral reefs are idyllic places
for shes to live, but dwelling in these shallow waters
means they often experience extreme water ows gener-
ated from waves, Dr. Fulton says.
Its almost like living with constant winds from a
cycloneyou can just imagine what itd be like to try and
nd food and get home in that sort of weather!
Having a smart swimming technique ensures that
these small reef sh have an evolutionary advantage in
the marine environment, Dr. Fulton says.
Fishes use up to half of their energy on swimming.
So if they can save even just a fraction of this, they can
spend it on growing bigger, holding larger territories, and
producing more offspring, he says.
Just imagine if you could save 40 percent on the
gasoline bill for your carhow good would it be to spend
that spare cash on other things?
We know that shes with these wing-like ns domi-
nate shallow reefs around the world, and in some cases
they can be about 10 times more abundant than shes
with paddle-shaped ns.
Fulton, Christopher J., Jacob L. Johansen, and John F.
Stefensen. 2013. Energetic extremes in aquatic locomotion by
coral reef fshes. PLoS ONE 8 (1): e54033. doi: 10.1371/journal.
From materials released by the ARC Centre for Excellence: Coral
Reef Studies, James Cook University, Townsville, Australia.
Chinas corals facing wicked problem
Chinas coral reefs have suffered a devastating 80 percent
decline in recent decades, driven mainly by the countrys
unrestrained economic development, according to a new
international scientic study.
The rst comprehensive survey of the state of corals
along mainland China and in the South China Sea re-
ports a grim picture of decline, degradation, and destruc-
tion resulting from coastal development, pollution, and
Professor Terry Hughes and Matthew Young of the
Australian Research Council Centre of Excellence for
Coral Reef Studies and James Cook University, and Dr.
The Wait Is Over
Once upon a time the way to
start a marine tank involved
introducing a few hardy fishes
and waiting until the tank
cycled. The wait to establish
an active biological filter often
lasted one to two months!
Nowadays the startup of a
marine aquarium is so much
simpler, and faster too!
Marine contains
cultured naturally occuring
microbes that rapidly start the
biological filtration process. Use
it to start the nitrification cycle in
new aquariums or to enhance
nitrification and denitrification in
heavily stocked aquariums.
What are you waiting for?
Two Little Fishies Inc.
1007 Park Centre Blvd.
Miami Gardens, FL 33169 USA
Hui Huang of the South China
Sea Institute of Oceanology,
Chinese Academy of Sciences,
published a new study in the
prestigious journal Conserva-
tion Biology that describes the
situation as a wicked prob-
lemmeaning it has no easy
A wicked problem is one
that is very hard to solve with-
out having a whole lot of other
foreseen and unforeseen con-
sequences to people and indus-
tries and to the environment
itself, Prof. Hughes explains.
Chinas ongoing economic
expansion has exacerbated many wicked environmen-
tal problems, including widespread habitat loss due to
coastal development, unsustainable levels of shing, and
pollution, the report states.
We found that coral abundance has declined by at
least 80 percent over the past 30 years on coastal fringing
reefs along the Chinese mainland and adjoining Hainan
Island. On offshore atolls and archipelagos claimed by
six countries in the South China Sea, coral cover has de-
clined from an average of over 60 percent to around 20
percent within the past 1015 years.
So far, climate change has affected these reefs far
less than coastal development, pollution, overshing,
and destructive shing practices. Ironically, these wide-
spread declines in the condition of reefs are unfolding
as Chinas research and reef-management capacity are
rapidly expanding.
The corals of the South China Sea region cover an
area of 11.5 square miles (30,000 km
), have high con-
servation value, and support the livelihoods of tens of
A fsherman tries his luck in the South China Sea
with the Hong Kong skyline across the bay.
thousands of shers. The fact that some reefs are claimed
by several different countries makes conservation and
management particularly difcult.
Typically, when a coral reef degrades it is taken over
by seaweedsand from there, experience has shown, it
is very hard to return it to its natural coral cover. The
window of opportunity to recover the reefs of the South
China Sea is closing rapidly, given the state of degrada-
tion revealed in this study, Prof. Hughes says.
The scientists conclude that the loss of coral cover
in the South China Sea, as elsewhere, is due mainly to a
failure of governance on the part of the nations respon-
sible for the marine environment.
China and other countries in the region have recent-
ly established a number of marine parks, but
they are too small and too far apart to pre-
vent the decline in coral cover, Hughes notes.
Governing wicked problems becomes
more challenging as they increase in extent
from local to regional or global scales, partic-
ularly where institutions are weak or nonex-
istent, the scientists caution. Cases such as
the Spratly Islands, which are claimed by six
different countries, highlight the dilemma.
There is no quick x to a wicked prob-
lem as complex as securing a sustainable fu-
ture for coral reefs in China and the South
China Sea, they add.
We suggest that gov-
ernance of Chinas coastal
reefs can be improved by in-
creasing public awareness, by
legal and institutional reform
that promotes progressive
change, by providing nan-
cial support for training of reef scientists
and managers, by expanding monitoring
of coral reef status and dynamics, and by
enforcing existing regulations that pro-
tect reef ecosystems.
The scientists suggest that Chinas
centralized system of government is well
equipped to quickly rescue the regions
imperiled coral reefs in collaboration
with neighboring countriesbut this will
require innovative leadership and strong
public support.
Hughes, T.P., H. Huang, and M.A.L. Young.
2012. The Wicked Problem of Chinas
Disappearing Coral Reefs. Conserv Biol, doi:
Map of the South China Sea coral regions at:
Chinas corals facing wicked problem. http://
facing-wicked-problem. Posted December 27,
Tortugas marine reserve yields
more, larger sh
With mounting bad news about the
plight of Caribbean corals and reef areas,
a protected area at the remote, western-
most tip of the Florida Keys is becoming
a model for using human intervention
to help threatened marine environments
A new NOAA research report nds
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that both sh populations and commer-
cial and recreational anglers have ben-
eted from no-take protections in the
Tortugas Ecological Reserve, the crown
jewel of the Florida Keys National Ma-
rine Sanctuary, which was designated in
2001 and encompasses 151 square nau-
tical miles. The report evaluates the ef-
fect of the reserve on the living marine
resources of the region and the people
whose livelihoods are connected to them.
The reports analysis of long-term so-
cioeconomic and scientic information
documented the following results after
the ecological reserve was designated in
- Cvcrslcd sccics sucl as Black
and Red Grouper, Yellowtail Snapper,
and Mutton Snapper increased in pres-
ence, abundance, and size inside the re-
serve and throughout the region;
- Annual gatlcrings o sawning
Mutton Snapper, once thought to be
wiped out from overshing, began to re-
form inside the reserve;
- Commcrcial catclcs o rcc slcs
in the region increased, and continue to
do so; and
- Mo nancial losscs wcrc cxcri-
enced by regional commercial or recre-
ational shers.
To assess economic effects of the area
closure, social scientists from NOAAs
Ofce of National Marine Sanctuaries
and the University of Massachusetts ana-
lyzed catch landings and revenues from commercial sh-
ers (reef shes, shrimps, Spiny Lobsters, and King Mack-
erel) and surveyed recreational shing guides operating
within the Tortugas region before and for ve years after
reserve protection.
This research shows that marine reserves and eco-
nomically viable shing industries can coexist, said
Sean Morton, sanctuary superintendent. The health of
our economy is tied to the health of our oceans. They are
not mutually exclusive.
Key West commercial shery landings had an esti-
mated value of $56 million in 2011, up from $40 mil-
lion in 2001, according to NOAAs Fisheries of the Unit-
ed States reports. Ocean recreation and tourism support
Populations of commercially important
species like the Red Grouper, Epinephelus
morio, increased following creation of the
Tortugas Ecological Reserve in the Florida
Keys National Marine Sanctuary.
approximately 33,000 jobs in the Florida Keys.
The ndings in this report are good news for NOAA
management efforts to enhance sheries and other nat-
ural resources in the Florida Keys, said Holly Bamford,
Ph.D., NOAA assistant administrator for the National
Ocean Service. The results are equally important in
other areas where NOAA science provides support to
management decisions that are made to best utilize and
protect our natural resources.
The design of the Tortugas Ecological Reserve in-
volved extensive collaboration between commercial
and recreational shermen, divers, scientists, conser-
vationists, citizens, and resource managers. The reserve
is closed to all consumptive use, including shing and
anchoring, and a portion of it is open only to permitted
marine researchers.
The Florida Keys National Marine Sanctuary protects
2,900 square nautical miles of critical marine habitat,
including coral reef, hard bottom, seagrass meadows,
mangrove communities, and sand ats, as well as ship-
wrecks and maritime heritage resources. NOAA and the
state of Florida manage the sanctuary.
From materials released by NOAA. Report: An Integrated
Biogeographic Assessment of Reef Fish Populations and
Fisheries in Dry Tortugas: Efects of No-Take Reserves.
Online at http://ccma.nos.noaa.gov/ecosystems/coralreef/
Many Caribbean reefs dead
in the water
The ability of coral reefs to
maintain their structures and
continue to grow depends
on the balance between the
addition of new carbonate,
which is mostly produced by
corals themselves, and the
loss of carbonate through
various erosional processes.
Scientists have long
known that reef ecosystems
are in decline and that the
amount of live coral on reefs
is dwindling, but the paper,
published on January 29 in
Nature Communications, is
the rst evidence that these
ecological changes are now
also having an impact on the
growth potential of reefs.
Our estimates of current
rates of reef growth in the Ca-
ribbean are extremely alarming, says
professor Chris Perry of the University
of Exeter, who led the research. Our
study goes beyond examining how
much coral there is, to also look at the
delicate balance of biological factors
which determine whether coral reefs
will continue to grow or will erode.
Our ndings clearly show that recent
ecological declines are now suppress-
ing the growth potential of reefs in the
region, and that this will have major
implications for their ability to respond
positively to future sea level rises.
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It is most concerning that many coral reefs across the
Caribbean have seemingly lost their capacity to produce
enough carbonate to continue growing vertically, whilst
others are already at a threshold where they may start to
erode. At the moment there is limited evidence of large-
scale erosion or loss of actual reef structure, but clearly if
these trends continue, reef erosion looks far more likely.
Urgent action to improve management of reef habitats
and to limit global temperature increases is likely to be
critical to reduce further deterioration of reef habitat.
The team was funded by the Leverhulme Trust (UK),
through an International Network Grant. It included
scientists from James Cook University and The Univer-
sity of Queensland in Australia, The University of Auck-
land in New Zealand, Memorial University in Canada,
and the University of Maine in the U.S. They examined
rates of carbonate production across 19 reefs in the Ca-
ribbean countries of the Bahamas, Belize, Bonaire, and
Grand Cayman.
Declines in rates of carbonate production were es-
pecially evident in shallow water habitats, where many
fast-growing branching coral species have been lost. The
study compared modern day rates with those measured
in the region over approximately the last 7,000 years.
As stony corals disappear in the Caribbean, they are often
replaced by macroalgae and/or gorgonians. Coral reefs build
their structures by producing and accumulating calcium
carbonate, essential for their maintenance and continued
vertical growth capacity. An international research team has
discovered that the amount of new carbonate being added by
Caribbean coral reefs in four countries is now signifcantly below
rates measured over recent geological timescalesin some
habitats as much as 70 percent lower.

In key habitats around the Caribbean, the ndings suggested that in waters
around 16 feet (5 m) deep, reef growth rates are now reduced by 6070
percent compared to the regional averages taken from historical records. In
waters around 32 feet (10 m) deep, the rates are reduced by 25 percent.
The study also suggests that these key habitats must have at least 10 per-
cent living coral cover to maintain their current structures. The amount of
cover varies between sites, but some are already below this threshold and are
therefore at risk of starting to erode. Given that previous studies have shown
that coral cover on reefs in the Caribbean has declined by an average of 80
percent since the 1970s, this raises alarms for the future state of reefs in
the region. These changes have been driven by human disturbance, disease,
and rising sea temperatures, and are only expected to intensify as a result of
future climate change.
From materials released by the University of Exeter, January 12, 2013.
A crab biology fest
Scientists who are willing to eat what
they study may represent a distinct
minority, but Judith S. Weis, a ma-
rine biologist at Rutgers University,
is equally happy envisioning crabs as
they evolved from lobsters during the
Jurassic period, describing how com-
mensal crabs that live with Acropora
and Pocillopora spp. stony corals in-
crease the sceleractinians growth rate
and clean potentially damaging sedi-
ments from their hosts, and even fol-
lowing the beautiful swimming portu-
nid crabs from the waters of Chesapeake
Bay to platters of Maryland crab cakes
with remoulade sauce.
Excerpt: In Belize, the land hermits,
Coenibita clypeatus, have organized shell
exchanges, in what is called a synchro-
nous vacancy chain by Randi Rotjan and
colleagues from Tufts University, who dis-
covered this unique behavior. When a large,
empty shell becomes available, many crabs
gather around it, which can take hours. As
they gather, the crabs arrange themselves
into a line of decreasing size, starting with
the largest crab holding onto the empty shell. As though choreographed, the
crabs begin shell-swapping one after the other, a smaller crab climbing into a
new shell right after it is vacated by the slightly larger crab ahead of it. What
makes the synchronous chain possible is that smaller crabs linger near a too-
large shell, perhaps attracting others, waiting until a bigger crab comes along,
which increases their chances of getting a good-tting hand-me-down.
Here is a highly readable book with a good balance of science and biology
trivia, which would make a valuable addition to any beach-house library or
serious aquarists bookshelf. It is perhaps best consumed in short doses, but
the reader is unlikely ever to think of the worlds 7,000 species of crabs in the
same ways again.
Walking Sideways:
The Remarkable World of Crabs
by Judith S. Weis
2012, Cornell University Press,
Ithaca, New York
256 pages, $29.95
False Coris Wrasses
Genus Pseudocoris
There are a number of wrasse groups that specialize in
feeding on zooplankton. These include two genera cov-
eted by aquarists, the fairy wrasses (Cirrhilabrus) and the
asher wrasses (Paracheilinus).
The members of the genus Pseudocoris also ll this
trophic niche. These shes spend much of their time
swimming in the water column (in some cases up to 33
ft. [10 m] over the seaoor), where they pick off passing
planktors. They also tend to occur in small- to medium-
sized shoals and often form mixed groups with other
zooplankton-feeding shes.
One clue to their open-water lifestyle is their lunate
caudal n. A tail like this is indicative of speed and is
a characteristic shared by many small open-water feed-
ers. These sh are torpedo-shapedanother physical clue
that they are speedsters. Their shoaling behavior is also
an adaptation to being more vulnerable high above the
reef. At least some species are known to refuge under the
sand at night.
While these wrasses are not commonly encoun-
tered in the aquarium store, the ve species in the ge-
nus make interesting introductions to the moderate- to
large-sized home aquarium. Not only are they active
shes that will need plenty of swimming space, but
some are also very colorful and will spend most of their
time in full view, dashing about in the water column. A
tank of at least 135 gallons is a minimum requirement
for a small group.
When rst acquired, the false corises may spend
much of their time buried in the substrate. It is important
Text & i mages by SCOTT W. MI CHAEL
Pseudocoris heteroptera, Torpedo False Coris Wrasse,
terminal male: built for speed, to capture zooplankton
in midwater.
to get their biological clocks reset to adjust to the new light-dark regime in
your aquarium. I have had individuals who always come out at night and
stay hidden during the daylight hours. In time, they usually will recover from
their jet lag. If you nd your Pseudocoris do not start to adjust after a week,
you may want to try and encourage them to come out of the sand by gently
probing the sand with your nger or a piece of rigid airline tubing. (Make
sure you do this gently, as you dont want to injure your labrid charges.) Some
aquarists place newly acquired wrasse species that bury in the sand at night
in a quarantine tank that lacks a sand bed, so the sh are forced to adapt to a
new light-dark cycle before being placed in a tank with sand to bury in.
The Pseudocoris tend not to be overly aggressive, although males may
pester other zooplankton-feeding labrids. Keep only one male per tank. Ju-
veniles and females can be housed in small groups and may do best if kept
in shoals or with other zooplankton-feeding wrasses (e.g., Paracheilinus).
That said, I have had females that were picked on by larger fairy wrasses
(Cirrhilabrus spp.). These wrasses (especially males or females that are being
harassed by more bellicose neighbors) will jump out of open aquariums. If
harassed by tankmates they typically hide incessantly and will not survive.
Pseudocoris yamashiroi, Redspot False Coris Wrasse,
initial phase: a classic zooplankton feeder and a
great reef aquarium species.
Pseudocoris yamashiroi, Redspot False Coris
Wrasse, terminal phase male: an unusual
variant from the Maldives, Indian Ocean.
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color up and grow like gangbusters.
Thats why we created AcroPower, our
new formula that supplies amino acids
that corals need to build their skeletal
architecture. Like other cnidaria, corals
have a special ability to uptake
dissolved amino acids across their
entire surface. Closed system
aquariums with protein skimming and
other ULN (ultra-low nutrient) filtration
methods deplete amino acids that are
vitally important for coral health. Corals
become more colorful within days due
to the extension of the growing margins
when AcroPower is used.
Poly-Filter has rescued tens of thousands of freshwater, brackish, saltwater and reef aquaria during the past
Thirty-Five years. During December 2011 alone we received ten emails, from experienced reef aquarists, all
asking about the blue coloration adsorbed into Poly-Filter. These long term, reef aquarists had not been
previously using Poly-Filter. Instead they had chosen: activated carbon, activated carbon and ion exchange
resins, macro reticulated styrene adsorbents and other sorbent media. However, when problems with the
corals developed each had decided these other filtration products didnt work. Maybe a Poly-Filter would
solve the problem?
The unique, Patented color change in Poly-Filter appeared to indicate
copper being adsorbed. How could this happen? Each of these
aquarists had been treating tap water using low pressure reverse
osmosis and mix bed deionization. We explained that if the
mixed bed D. I. resins failed copper would certainly enter the
aquaria. Could ten very experienced reef aquarists, all having
mix bed resins fail? It is certainly possible, low pressure reverse
osmosis will bypass sufficient copper that it would negatively
impact corals. However, we suspected additional copper and
other heavy metals, so inquiry was made about amount, frequency
and types of water treatments and/or coral additives being
used. Every reef aquarist had been dosing: calcium, magnesium,
strontium, iodine /iodide, carbonates and trace elements. Would
copper, iron, lead be trace contaminates found in these additives?
It is a scientific fact that lead is the major contaminate in all
calcium compounds! Iron is another almost a universal contaminate.
Copper may have entered aquaria as a trace contaminate found
in the trace elements or other coral additives? However when
Poly-Filter adsorbed the copper it would also adsorbed: iron,
lead, excess heavy metals, dissolved organics, phosphates, volatile
organic chemicals, pesticides and any biotoxins. Water quality was
corrected, aquarists reduced the amount of additives and the clarity
of their aquaria greatly increased. Coral health and growth problems
solved! Poly-Filter adsorbed 31.97% of 718.95 micrograms per liter
ionic copper @ 14.307 liters per minute within seventeen seconds.
Activated carbon/resins (283 grams) adsorbed only 20% of the 718.95
micrograms per liter of copper but needed
30 seconds. Zeolite resin adsorbed 14.5% of the 718.95 micrograms per liter of copper within
17 seconds. Poly-Filter also adsorbed 61.8% of 211 micrograms per liter within 60 seconds at 14.307 liters
per minute out of saltwater. Poly-Filter adsorbed 46.4 % of 1570.2 micrograms per liter of chelated copper
@ 14.307 liters per minute within 14 minutes. A strong chelating resin (1000 ml.) could only adsorb 13.03 % of
1570 micrograms per liter @ 14.307 liters per minute within 14 minutes.
Feed them frequently, at least three times a
day. They greedy accept frozen mysid shrimp,
and nely minced table shrimp; smaller indi-
viduals will snap up frozen Cyclops.
The most common species in aquarium
stores is the Redspot False Coris (Pseudocoris
yamashiroi, Schmidt 1931). The male of this
species has a white ventrum and is gray, green,
and yellow on the upper third of the body with
scattered dark spots. Females are pink to gray
overall with silver or blue lines on the head
that can extend onto the body, especially in smaller individuals. It reaches a
length of just over 15 cm (5.9 in.).
Pseudocoris yamashiroi ranges from East Africa to Samoa, south to the
Kermadec Islands, and north to the Philippines. It is found over lagoon patch
reefs, in reef channels, on forefaces and slopes at depths of 7 to at least
83 feet (225 m). This species forms aggregations as juveniles and adults.
Groups of the latter are composed mainly of females with the occasional
male. It is a zooplankton feeder that ingests copepods and mysid shrimps. The
Redspot False Coris is a wonderful addition to the reef tank. Only one male
should be housed per tank and a male is best introduced with or after a group
of females. This species will leap from an open aquarium. Feed a meaty food
(e.g., chopped seafood, mysid shrimp) three or more times a day.
Bleekers False Coris (Pseudocoris bleekeri, Hubrecht 1876) is a lovely sh
that is known from Indonesia north to the Ryuku Islands. Female P. bleekeri
are green, while the male has a distinct bright yellow band on the side (this
is more conspicuous when the sh are feeding over the substrate). It reaches
a length of 5.9 inches (15.0 cm). This false coris is found over coastal sand
and rubble reefs and over coral pinnacles, usually in current-prone areas.
Pseudocoris yamashiroi,
Redspot False Coris Wrasse:
a group of initial phase fsh.
Members of this genus tend
to gather in clusters, which
ofers them some safety when
they feed away from cover.
One terminal phase male
typically rides herd over a
number of initial phase fsh.
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Reactor 150.
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dispersion plate, they force an even distribution of
water through the media. Mount them on the back of
the aquarium or below it. Each reactor includes a ball
valve for regulating flow, and flexible connection
fittings that rotate 180 degrees to allow a perfect
custom fit to your installation. The threaded lid design
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It feeds on zooplankton and often
forms mixed assemblages with other
zooplanktivores. When the male rises
into the water column to feed, the
yellow blotch appears on the side. Its
husbandry requirements are similar to
those of P. yamashiroi.
The Torpedo False Coris or Torpe-
do Wrasse (Pseudocoris heteroptera,
Bleeker 1857) is distributed from the
Seychelles to the Society Islands, north
to south Japan. It reaches a length of
6.3 inches (16 cm). The males exhibit
an attractive color pattern consisting
of a bluish gray head and anterior
body. The posterior section of body is green with dark
bars. Females are white to tan with a dark brown stripe
along the back and a thicker stripe running down the
center of the body. The anal n is often orange. The Tor-
pedo False Coris is a resident of outer reef crests and reef
faces, where it occurs at depths of 7 to 178 feet (254
m). It is often found over open bottoms composed of a
mixture of sand and rubble, with scattered coral heads.
Pseudocoris heteroptera lives in small groups comprised of
many initial phase sh and a single terminal phase male.
When initial phase sh move up in the water column to
feed, they often lose their stripes and become a mono-
chromatic gray, which makes them less conspicuous
to roving piscivores. Males exhibit rapid color changes
when interacting with rivals or potential mates. Hus-
bandry is similar to that for others in the genus.
Pseudocoris bleekeri, Bleekers False Coris
Wrasse, terminal phase male, Cebu.
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and invertebrates.
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and invertebrates.
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spp. in the
aquarium of C.
oanthids (also known as encrusting or colonial
anemones, sea mats, and button polyps) are
among the sessile invertebrates that were available in the
early days of the reef aquarium hobby. As long ago as the
1970s, Peter Wilkens, the late Swiss pioneer of the reef
aquarium hobby, was maintaining colonies of encrusting
anemones. Zoanthus and Protopalythoa polyps can still
be found in coral reef aquariums, but the enthusiasm of
aquarists for these colorful polyps has waxed and waned
over the past three decades.
article and images by Dani el Knop
Zoanthus sp.
In the 1990s, when it be-
came possible to maintain
small-polyp stony corals in
the aquarium, they grew
very popular, and interest in
zoanthids, which cover the
rockwork with an attractive
carpet, diminished. Dealers
shoved them into the back
corners of sales aquariums,
and aquarists exiled them
from their reef tanks to
make way for the coveted
stony corals and splendidly
colored Acropora speci-
mens. By the late 1990s,
few coral farmers would
have dreamed of deliberate-
ly propagating encrusting
These actinians became
even more unpopular when
articles about their toxicity
appeared in aquarium mag-
azines. Reports of palytoxin
contamination via skin in-
juries and warnings about mass proliferation in the reef
aquarium didnt help. Rolf Hebbinghaus, for example,
told me about a dramatic mass proliferation of a Zoan-
thus species in the huge semicircular reef aquarium at
the Lbbecke Aquazoo, and despite intensive efforts he
was unable to curb their spread. By the end of the mil-
lennium the passion for zoanthids had reached a nadir
in Europe. Encrusting anemones in the aquarium? Not
for me!
The trend in the opposite direction began in the
United States, where numerous aquarists longed for cer-
tain Zoanthus color morphs and were willing to pay a
small fortune for a tiny group of polyps on a piece of
coral rubble. Aquarists in North America tended their
orange, yellow, red, and blue Zoanthus jewels, swapped
photos, and produced small rocks with cuttings consist-
ing of three, four, or ve individual polyps, which ulti-
mately changed hands for prices that were being paid
for a rare, steel blue Acropora echinata in Germany at the
time. Its a mad world!
A few years later, reef aquarists in Europe began to
show renewed enthusiasm for these colorful actinians.
Gradually, an interest in encrusting anemones stopped
being something to be ashamed of among reef aquarium
hobby specialists. But anyone who is looking for a hus-
bandry challenge shouldnt be considering encrusting
anemones, as they are so robust and undemanding that
they are ideal for the beginner.
Unfortunately, the encrusting anemones of the order
Zoanthidea have for a long time been in a sort of sci-
entic Sleeping Beauty slumber. Today, very few details
of their systematics and phylogenetic relationships are
known, and what is known is often based on the former
taxonomic practice of classifying the animal kingdom on
the basis of characters of physical form (morphology),
without regard for actual phylogenetic relationships. Sci-
entic taxonomy is currently experiencing what is prob-
ably the greatest upheaval since its inception; genetic
research methods are allowing scientists to study and
reclassify the animal kingdom according to its phyloge-
netic relationships, and many earlier taxonomic classi-
cations have proved to be incorrect.
Knowledge of the encrusting anemones is also grow-
ing due to this development, as specialists like Dr. James
D. Reimer in Japan are studying them intensively. Using
comparative DNA studies, they are determining phy-
logenetic relationships and producing cladograms that
show lines of descent. Many genera that are well known
and popular in the aquarium hobby, such as Palythoa
and Protopalythoa, will also undergo changes within the
framework of this study. In Professor Reimers opinion,
these genera include several species that arent closely re-
lated at all. Before genetic research methods were avail-
able, they were grouped together on the basis of their
similar body structures and anatomical details. But now
that phylogenetic relationships at genus level can be de-
termined with greater accuracy, in a few years the reef
aquarium hobby will have access to new information
about these attractive actinians that have been kept in
aquariums for over 30 years.
Protopalythoa sp.
Encrusting Anemones
Things worth knowing about
ncrusting anemonesattractive to look at, undemand-
ing, able to look after themselves, and very toxic in
some casesprovoke a range of reactions among reef
aquarists, from affection and passion to revulsion.
The majority of zoanthid species are very smalltheir
mouth-discs are rarely more than .4 inch (10 mm) in diameter.
They do not produce a solid calcareous skeleton; instead, their
polyps exude a layer of mucus to which foreign bodies, mainly
ne sediment particles, adhere. This mixture of sediment and
mucus secretion solidies and is overgrown by the outer skin.
In this way they develop an inner auxiliary skeleton that can
contract to a greater or lesser degree, depending on the genus;
some can only close their mouth discs and contract mildly
(for example, Protopalythoa species), while others can shrink
to almost unrecognizable little lumps of tissue that are barely
visible on the substrate (for example, the Yellow Encrusting
Zoanthids are rapid secondary colonizers that immediately
occupy any place that becomes vacant on the reef. Many of their
colonies occur in the tidal zone between the high and low water
marks, a habitat to which most other creatures cannot adapt.
They protect themselves from dehydration and excessive solar
radiation by releasing large amounts of mucus when exposed.
Zoanthus sp.
article and images by Dani el Knop
Palytoxin is a highly toxic natural substance whose
provenance has not yet been completely explained. It
is known to be present in certain dinoagellates (Os-
treopsis siamensis), and this has led some scientists to
assume that encrusting anemones extract it from cap-
tured plankton and use it to protect themselves against
predation. However, this assumption is contradicted
by the fact that not all encrusting anemones that pos-
sess palytoxin practice intensive plankton capture, and
the poison is also present in encrusting anemones that
have grown in the aquarium without ever having the
opportunity to capture the dinoagellate Ostreopsis
siamensis. In light of this, some theorize that the toxin
is manufactured in zoanthids by bacteria.
Palytoxin has a powerful cytolytic (cell-dissolving)
effect and is one of the most powerful animal-derived
poisons known. Contact with the animals and their
mucous secretion is harmless as long as the skin is un-
broken. But if the poison gets into the skin through a
puncture wound or cut, it triggers a
severe inammation leading to ne-
crosis of the surrounding tissue (cy-
tolytic effect). Even when intact, the
cornea of the eye is very delicate; if
the poison accidentally gets in the
eyes, it can cause massive inamma-
tion and damage to the sight; cloud-
ing of the cornea can even lead to
transitory loss of vision.
In several cases, severe, allergy-
like symptoms of an anaphylactic
nature have been described follow-
ing the inhalation of aerosol (air
containing tiny drops of water) con-
taining palytoxin from Protopalythoa
heliodiscus, necessitating a hospital
stay of several weeks for the victim.
The indigenous peoples of Hawaii
used palytoxin to make poisoned ar-
rows. The toxic effect, which sets
in rapidly and can lead to a painful
death within a fairly short time, comes about through
the destruction of cell-specic ion-transport systems.
The existing concentration ratios for potassium and
sodium ions go out of balance, which has a dramatic
effect on the cells and the entire organism as a result of
disturbed osmotic pressure, altered membrane poten-
tial of the cells, and other effects. In experiments with
mice, as little as 100 nanograms (ng, one billionth of
a gram) per kilogram of body weight proved fatal for
50 percent of the experimental animals (LD50), and a
carcinogenic effect was demonstrated.
We marine aquarists should wash our hands thor-
oughly with soap after any contact with encrusting
anemones, including Zoanthus species. When working
on encrusting anemone colonies out of water (for ex-
ample, when removing the polyps from the substrate)
it is prudent to wear protective gloves, goggles, and a
face mask to avoid inhalation of palytoxin-containing
The majority of species of zoanthids produce exten-
sive mats or colonies consisting of numerous individual
specimens. The very few species that live as solitary pol-
yps include the members of the genus Sphenopus. Three
different habits of growth can be distinguished in the
colonial-living species:
1. The individual polyps exhibit solitary growth,
without being connected to their neighbors;
2. all individuals in the colony are linked together
via basal runners;
3. all polyps are embedded in a thick shared coenen-
chyme, and only the mouth discs are visible, not
the body columns.
The mouth disc bears numerous tentacles grouped in
one or two rings around the centrally positioned mouth
opening. Many species are particle-feeders, and most live
in symbiosis with unicellular zooxanthellae, with symbi-
otic algae that also lend them their dark color. Many spe-
cies protect themselves against being eaten by predators
by means of a poison called palytoxin, one of the most
toxic of all known natural substances, which was named
for the encrusting anemone genus Palythoa, in which
Palytoxin precautions
When touched,
heliodiscus releases
large amounts
of a clear mucus
secretion with an
extremely high
palytoxin content.
Ornate Ghost Pipefsh,
Solenostomus paradoxus,
pair (larger female, left).
it was rst discovered. There is no
precise information about the ori-
gins of this poison, but scientists
currently believe it is produced by
bacteria that live in symbiosis with
the polyps. The poison is most con-
centrated when the polyps produce
Some 300 zoanthid species are known from all the
seas. However, at present it is possible to classify them
precisely in only a few cases, as the systematics of en-
crusting anemones have not been well studied. Genetic
study is sometimes necessary even for differentiation to
species level, so that specialty taxonomists are unable to
assign many encrusting anemones to a particular spe-
cies simply on the basis of external appearance. Making
matters even more difcult is the new trend in scientic
taxonomy away from categorization on morphological
characters and toward division on the basis of actual
and demonstrable phylogenetic relationships. Systematic
classication is being re-examined in practically all areas
of marine biology, and much will be changed as a result.
Some genera of encrusting anemones known today will
no longer exist in a few years, because they have been split
up and their members assigned to other genera.
The encrusting anemones of the order Zoanthidea
are divided into two groups. The important character for
this division is differences in the fth mesenterium. A
mesenterium is a sail-like partition in the interior of a
polyp, extending from the center to the outer wall. Those
with a complete fth mesenterium are termed Mac-
rocnemina, while those with an incomplete fth mes-
enterium are Brachycnemina. This distinction is fairly
meaningless for the aquarist, but an awareness of these
two suborders will help to summarize the phylogenetic
The suborder Brachycnemina is by far the most impor-
tant for the reef aquarist, as it contains the two families
Sphenopidae (genera Palythoa and Protopalythoa) and
Zoanthidae (genus Zoanthus). The suborder Macrocne-
mina contains mainly families with fewer species suit-
able for the aquarium: Epizoanthidae, Hydrozoanthidae,
and Parazoanthidae. But it isnt irrelevant for the ma-
rine aquarium hobby: besides the numerous Parazoan-
thus species that are sometimes imported with sponges,
it includes an encrusting anemone species that is so far
scientically undescribed but regularly sold in the aquar-
ium trade: the Yellow Encrusting Anemone, or Colonial
Yellow Polyps. Its original description is imminent and
will place it in the genus Terrazoanthus.
If the tentacles are
densely packed, every
second one curves
slightly toward the
center of the mouth
disc so that two rings of
tentacles result.
The phylogenetic relationships of the individual
species of encrusting anemones are currently
being studied.
The Zoanthus species
belong to the suborder
The family Zoanthidae contains encrusting anemone
genera that are frequently encountered in the aquarium
trade. This applies above all to Zoanthus. Specimens of
Isaurus and Acrozoanthus only occasionally appear in the
aquarium trade, and as a rule they prove to be very prob-
lematical in their husbandry.
Genus Zoanthus
Encrusting anemones of the genus Zoanthus live world-
wide on tropical and subtropical reefs and in coastal
zones. They live throughout the entire strongly lit zone,
even in very shallow waterincluding in the tidal zone
and are extraordinarily robust in the reef aquarium. As
a rule they form relatively large, dense groups of polyps
that completely cover the substrate, sometimes blanket-
ing enormous expanses of rock.
There are 22 valid species, and a number of others are
still being investigated (species inquirenda). However,
the differentiation of the species doesnt depend solely on
coloration, so it is virtually impossible in the aquarium
hobby. In science it is facilitated by DNA comparison.
One of the best-known species is Z. sociatus. It is found
mainly in the Caribbean, but small populations occur in
parts of the western Pacic, for example south of Japan.
Most Zoanthus species are usually labeled as Zoanthus so-
ciatus in the marine aquarium hobby, but this is incorrect
unless they are members of this Caribbean species.
Zoanthus species form a thin basal layer on the sub-
strate, from which the polyps sprout. The polyp consists
of a body column that broadens at the top and forms a
mouth disc bordered by a ring of tentacles. If the ten-
tacles are pumped up by the internal hydraulic pressure
of the polyps to such an extent that they become very
crowded, every second tentacle extends forward, so that
there are two rings of tentacles, one behind the other.
Zoanthus sp.
Zoanthus sp.
Two Zoanthus spp.
The centrally positioned mouth opening is often a con-
trasting color. It is very interesting to maintain color
morphs that have uorescent markings. Blue light in-
duces green uorescence in these, particularly visible un-
der completely blue lighting.
Genus Isaurus
The genus Isaurus contains 10 species, only one of which
occasionally turns up in the aquarium trade: the azoo-
xanthellate I. tuberculatus. The body columns are
considerably longer than those of the externally
similar genus Protopalythoa, and they normally open
their polyps only at night.
The species isnt dependent on light; it lives by
capturing food. Its brown coloration leads many
aquarists to assume that it has symbiotic algae, but
that is not the case, so it must be fed intensively.
Unfortunately, knowledge of its azooxanthellate way
of life isnt widespread in the trade, so these polyps
are almost never regularly fed during the export and
trade stages. Consequently, when the polyps reach
the retailers sales aquarium they are usually already
so weakened and degenerated that they can no lon-
ger be established successfully in the aquarium. In
order to accomplish this you must feed them directly
several times a day.
Genus Acrozoanthus
The genus Acrozoanthus contains only the species A. aus-
traliae. The polyps look almost like little sea anemones,
but they belong to the Zoanthidae encrusting anemones.
As a rule they settle on exible worm tubes. These azoo-
xanthellate encrusting anemones sometimes turn up in
the aquarium trade, but they are so sensitive to trans-
portation that it is rarely possible to establish them in
the aquarium.
Isaurus tuberculatus, with open
and closed (below) polyps.
The family Sphenopidae contains two further encrusting
anemone genera that are of importance for the aquari-
um hobby: Protopalythoa and Palythoa. The third genus,
named Sphenopus, plays practically no role in the aquar-
ium hobby, as it contains three species that live not in
groups but as individual zooids.
Genus Protopalythoa
After Zoanthus, the genus Protopalythoa is the most com-
mon in the aquarium hobby. But at present it is a sort of
taxonomic catch-all for several polyphyletic encrusting
anemone species (those that dont represent any shared
phylogeny). In the future the DNA of these species will
be examined in order to ascertain their actual relation-
ships, so the genus may be split into several genera.
Encrusting anemones of the genus Protopalythoa live
throughout the bright-light
zone on the reef to a depth
of around 33 feet (10 m),
also occurring in very shal-
low water, and are extremely
robust. As a rule they grow
in small groups. They can
be distinguished from the
similar genus Zoanthus by
their larger-diameter mouth
discs; also, their tentacles
have pointed tips.
They are different from
the genus Palythoa in that
the polyps do not have a
shared body mass, but are connected with one another
only by a thin basal layer attached to the substrate. The
largest species is thought to be Protopalythoa grandis,
which attains a mouth-disc diameter of up to 2.4 inches
(6 cm); there are probably at least two species that grow
this large, but only one is scientically described.
Protopalythoa heliodiscus
Protopalythoa heliodiscus is a somewhat unusual encrust-
ing anemone species. It has a similar appearance to the
green Protopalythoa known in the aquarium hobby, but
it is gray. Its body column is consider-
ably softer, as it accumulates less sed-
iment in the wall. For this reason its
polyps can also contract much more
than those of all the other Proto-
palythoa species. It also releases large
amounts of a clear mucous secretion.
Be very careful with this secretion: it
contains a high concentration of pa-
This species is more able than
other encrusting anemones to take
in planktonic food, as in the wild it
lives in twilit zones where it compen-
sates for the lack of light by captur-
ing plankton. The strong lighting of a
reef aquarium is unnatural for it and
sometimes triggers a dramatic mass
proliferation that is difcult to curb
because of the high palytoxin con-
Protopalythoa heliodiscus is the
only encrusting anemone for which
aquarium maintenance is inadvisable
for safety reasons. There is a danger of
mass proliferation and an accompa-
nying high production of a toxin that
can also be deadly to humans. If this
species is present in the aquarium,
the population should be monitored
Acrozoanthus australiae,
Polyp Trees or Stick Polyps
and Zoanthus
mouth discs for
to ensure that it doesnt develop into a difcult-to-con-
trol plague. Great care is advised when it comes to me-
chanical control, as the secretion that is released on con-
tact, and in particular in the event of injury, is very toxic.
Genus Palythoa
Like Protopalythoa, the genus Palythoa is a taxonomic
catch-all for several zoanthid species that do not share
a common ancestry. Some notable taxonomists even re-
gard Protopalythoa as invalid and place its members in
the genus Palythoa, as was usual until a few years ago.
About 90 species are known, but they may include mul-
tiple descriptions. This is an area where phylogenetic re-
lationships need to be claried with the aid of genetic
studies in the years to come.
Encrusting anemones of the genus Palythoa live
throughout the strong-light zone of tropical coral reefs
to a depth of around 33 feet (10 m), also occurring in
very shallow water, and are ex-
tremely robust. As a rule they live
in small groups of polyps. They
differ from the genus Protopaly-
thoa in that the polyps are located
in a thick, eshy-looking, shared
body mass, which, however, only
becomes visible in many species when the polyps are
partially closed, as the open mouth discs cover it. In
Protopalythoa species, by contrast, the polyps grow
from a thin basal layer and exhibit individual, free-
standing body columns. Palythoa polyps contain the
highly poisonous palytoxin named after their genus.
The suborder Macrocnemina contains families that
are less important for the aquarium hobby, but
some genera turn up now and then in the trade, one fair-
ly regularly. Parazoanthus species are occasionally seen
for sale along with their host sponges. Most members of
this suborder are, however, signicantly more difcult to
keep in the aquarium than those previously described.
One exception is a scientically undescribed species that
is assigned to the genus Terrazoanthus: the Yellow En-
crusting Anemone (see page 40).
The family Parazoanthidae contains encrusting anemo-
nes that live commensally on other invertebrates. Only
the genus Parazoanthus, which groups together the species
that live with sponges, is interesting from an aquarium-
hobby viewpoint. It is conceivable that in many cases this
is a reciprocal relationship from which both partners de-
rive benet: the sponges prot from the defensive capa-
bility of the encrusting anemones, while the sponge offers
Protopalythoa heliodiscus spreading
invasively among other encrusting
anemones in the aquarium.
Palythoa. All of
the polyps are
embedded in
a thick shared
the anemones a substrate into which they can withdraw
and be protected by some of the secondary metabolites
of the sponge that are poisonous to predators. But a lot
of detailed taxonomic work is required in this family as
well; even many of the species that occasionally turn up
in the aquarium hobby are not yet scientically described.
Genus Parazoanthus
This genus currently contains 10 species, only one of
which lives in the Pacic: P. darwini from the Galapagos
Islands. All the other species originate from the Atlantic.
However, some known species are still awaiting scientic
description. Such undescribed zoanthids sometimes also
turn up in the aquarium trade.
Encrusting anemones of the genus Parazoanthus live
on and/or in sponges. Sometimes the host sponge can be
seen only when the polyps are open, if at all; only when
you touch the polyps so that they close does the sponge
come into view. Parazoanthus species have no symbiotic
algae, but live solely by capturing food. This makes their
aquarium maintenance difcult. In addition, because the
host sponge can be very sensitive to transportation, often
only the polyps are alive on arrival or after the rst days in
the wholesalers tank, while the sponge is disintegrating.
This explains why Parazoanthus species are rarely encoun-
tered in the hobby. If they do make it into a reef aquarium
unscathed, they can be established only via special care in
a species tank with suitable planktonic food, and the food
requirements of both the encrusting anemones and the
sponges must be taken into account.
The family Epizoanthidae contains encrusting anemones
that attach themselves to abandoned tubeworm casings.
This family is intrinsically unimportant for the aquarium
Two scientifcally
species on their
host sponges.
hobby, but occasionally polyps of the genus Epizoanthus,
with their worm tubes, turn up for sale, though so far
they havent been maintained successfully.
Genus Epizoanthus
The genus Epizoanthus contains around 85 species that
live on sessile invertebrates, analogous to the members
of the genus Parazoanthus. One example is Epizoanthus
illoricatus, which lives on empty worm tubes and extends
the latter in a characteristic zigzag form. Its polyps are
either black or pale yellow.
The encrusting anemone family Hydrozoanthidae at
present contains only two genera: Hydrozoanthus (three
species) and Terrazoanthus (one species). This family
would be of no signicance for the aquarium hobby, were
it not for the fact that the Yellow Encrusting Anemone,
known in the hobby but not yet scientically described,
is closely related to its members.
Genus Terrazoanthus
The genus Terrazoanthus, erected in 2010 by
Reimer & Fuji, currently contains only Terrazo-
anthus onoi, which plays no role in the aquar-
ium hobby. However, the scientic description
of a further species is in preparation, the one
known in the aquarium hobby as the Yellow
Encrusting Anemone. This little anemone from
the Indo-Pacic, which has turned up regularly
in the aquarium trade for more than two de-
cades, will purportedly be placed in the genus
Terrazoanthus. The names usually used for this
species in the aquarium hobbyParazoanthus
axinellae and Para-
zoanthus gracilisare
incorrect, as they de-
note Mediterranean
species that do exist
but have nothing to do with this
tropical encrusting anemone.
The Yellow Encrusting Anem-
one forms small groups of pol-
yps in which the individuals are
densely packed together. An indi-
vidual polyp consists of a slender
body column that broadens at
the top and forms a mouth disc,
bordered externally by a ring of
tentacles. The entire body is yel-
low to gray-yellow, depending
on the light intensity. The color
is inuenced by the symbiotic
algae, whose population density
depends on the strength of the
light. Under bright lighting the
coloration becomes darker.
These encrusting anemones cover rocky substrates
with a yellow carpet of polyps. Given adequate lighting
and generous feeding, this layer will become so dense
that the substrate is no longer visible. They live through-
out the strongly lit zone and are very hardy. When dis-
turbed, they contract into inconspicuous little heaps of
tissue. They must be regularly fed with planktonic food
in the aquarium in order to proliferate well.
The genus Neozoanthus was erected in 1972 by Herbert
and, to date, contains only one species, N. tularensis. Ad-
ditional species do exist, but they havent been described.
Neozoanthus species live in the Pacic and can arrive in
the aquarium with living rock or coral substrate rocks.
The genus Microzoanthus was erected in 2011 by Fuji &
Reimer and so far contains only the two species M. oc-
cultus and M. kagerou.
Epizoanthus illoricatus encrusting
tubes secreted by a sedentary
species of polychaete worm.
Right: The Yellow
Encrusting Anemone, a
scientifcally undescribed
Terrazoanthus species.
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46 CORAL 46 46 46 COR CORAL AL
aquarium husbandry and propagation
article by CI aude Schuhmacher - images by Dani el Knop
n 1985 I was importing and selling not only the very popular green (and poison-
ous) Protopalythoa but also the Yellow Encrusting Anemone that is still familiar
to everyone today but has not been scientically described. In those days we never
dreamed of the unusual and brightly colored color variants with imaginative names
that are easy to nd now, such as the blue Polar Ice Cap and the red Bali Cracker
(both from Indonesia), and the Cherry Mint from Taiwan. The Real Watermelon
and the Yellow Watermelon from Vietnam are among my personal favorites.
Because aquarists could only dream of such creatures until after the turn of the
millennium, it is no wonder that the zoanthids available in the pioneering days of the
marine aquarium hobby, which played an important role back then, were pushed into
the shadows by the arrival of the stony corals.
Back then, I was working for a marine wholesaler, and in retrospect I can only
describe our dealings with these creatures as reckless and uninformed. Of course we
Zoanthids, or sea
mats, were among
the rst sessile
invertebrates to be
imported for the
marine aquarium
hobby. In earlier
decades they were
invariably available
in only two or three
color morphs, usually
brownish Zoanthus
with orange or
yellow centers or
green Protopalythoa.
These days, there is
a regular panoply of
very brightly colored
variants to choose
Left: Encrusting
anemones can be kept
with other fuorescent
actinians to create
attractive populations.
Right, top: Encrusting
anemones of the
genus Zoanthus are
undemanding aquarium
Right, bottom: It
is easy to keep
encrusting anemones
with numerous other
were already aware that encrusting anemones were toxic,
but nevertheless we divided large clumps to make them
more saleable, usually in a rather primitive way with
a sharp knife or with hammer and chisel. The precise
techniques for taking cuttings that are normal nowadays
were unknown to us. Astonishingly, as far as I know,
there were none of the poisonings that seem to occur
increasingly frequently nowadays. Perhaps it was simply
the luck of the ignorant.
Aquarists and the aquarium trade began to take note of
particularly colorful zoanthids around 1995. More and
more colorful Zoanthus were becoming available, usually
as squatters on living rock or the substrate rocks of
soft corals. American dealers and aquarists, especially,
quickly discovered the potential of these colorful trea-
sures from Indonesia, and the fad soon spread to Europe.
Most of the fanciful names for encrusting anemone col-
or variants originated in the United States.
But the Zoanthus trend didnt really arrive in Europe
until 2003, when people began to import gaudy speci-
mens from the U.S., or directly from their native lands,
and sell sections of colonies as cuttings. Between 1998
and 2002 I was able to propagate these little jewels on
coral farms in the Philippines and pass my knowledge
on to other farmers.
After about 2005, the trend toward the keeping of color-
ful encrusting anemones slowly but surely became a veri-
table boom. So-called ultra variants of Zoanthus and
Palythoa reached central Europe via Hong Kong, Taiwan,
and Vietnam. But because these variants were difcult
to obtain and in great demand, some aquarists sought to
import established cuttings from the U.S., where these
animals were already more widespread.
I still remember bringing the rst Real Watermelon
Zoa back from the MACNA conference in a cookie tin.
The vendor was not a little surprised when I bought up
his entire stock. In those days one had to make an effort
to obtain particular specimens, although a number of
variants were already available direct from Fiji, Tonga,
the Caribbean, and Kenya as well.
Because encrusting anemones are basically easy to
maintain, dont require expensive equipment, and thrive
Encrusting anemones of the genus Zoanthus.
in aquariums that are small or have too
little light for SPS corals, for example, their
success story is hardly surprising. Increased
availability from online propagators meant
that they rapidly achieved immense popu-
larity. Buyers kept demanding more strik-
ing colors, and many an aquarist developed
a real passion for collecting these animals.
In 2005 I began to culture Zoanthus on
a larger scale and tried to develop standard
procedures for optimal results with the aid
of special tools and thorough experiments
that included targeted feeding. I mainly
used methods from the United States,
which I adapted or improved depending
on the species being bred. In the course of
this work I established that in no way are
all variants of Zoanthus similarly easy to
maintain. In particular, specimens import-
ed from the subtropics and Indonesia can
present difculties, especially with regard
to bacterial or parasitic diseasesa problem
that I will discuss in more detail.
Essentially, encrusting anemones are suit-
able for any reef aquariumvirtually re-
gardless of its size and technical equipment.
The more robust types are good for lling
spaces where other corals dont thrivelike
overow boxes and tubes, which can be al-
lowed to become covered with zoanthids,
and shady spots, for instance beneath large corals.
Keeping encrusting anemones with other aquarium
occupants is uncomplicated in
most cases. However, care must
be taken to make sure the crea-
tures are compatible; some LPS
or SPS, for example, can vigor-
ously sting Zoanthus, and them-
selves may fall victim to stinging
attacks by Palythoa species or
Yellow Encrusting Anemones.
The concept of zoanthids
for every reef aquarium also
has other limitations, at least
with certain ultra Zoanthus
from Japan or northern Taiwan.
These are often very sensitive
and exhibit problems during ac-
climatization to the conditions
usually found in reef aquari-
ums here, as they originate
from cooler waters. These types
should, if possible, be kept in
lower temperatures and rather muted lighting, insofar
as this is in keeping with the needs of other occupants.
Often posed, but difcult to answer, is the question of
optimal aquarium conditions for zoanthids. It has been
more or less established that stable water parameters
and low nutrient concentrations are essential for success
in maintenance and propagation. That being said, they
can also be kept in water with a higher nutrient load,
but this does have disadvantages: the extremely colorful
Zoanthus species develop their colors only at low nutri-
ent levels, and certain Protopalythoa species tend to the
dreaded mass proliferation if the nutrient supply is el-
evated. Over the years I have found the following water
parameters to be advisable:
pH: 8.28.4
Calcium: 400440 mg/L
Magnesium: 1,2501,350 mg/L
Carbonate hardness: at least 7.59dH (Zoanthus toler-
ate low alkalinity poorly)
Temperature: 7580F (2427C) (with the exception
of subtropical species)
Nitrate: up to 10 mg/L
Phosphate: up to 0.08 mg/L
Even though encrusting anemones will thrive un-
der various lighting conditions, there is a useful rule
of thumb for providing the correct illumination: very
brightly colored specimens should have moderate to
strong lighting. On the other hand, for dark-colored
specimens, weak, indirect lighting will produce better
results. This is particularly important for accentuating
the uorescent effects often present in such specimens.
Particularly attractive encrusting
anemonesthese are Zoanthus
specimenscan be attached to
substrates with cyanoacrylate
adhesive gel.
My experience shows that a blue-biased light spectrum
is best for this and can be achieved most easily and ef-
fectively with LED lighting.
Another important factor for the long-term health of
encrusting anemones is regular partial water changes us-
ing top-quality salts; if possible, do not add any carbon-
based substances such as biopolymers. Marine salts that
are heavily augmented with certain elements,
such as are increasingly the fashion for the main-
tenance of stony corals, should also be avoided.
Loss of brilliance in the colorsassuming this
actually represents a deciencyis far better rec-
tied deliberately with trace-element solutions.
In such cases, spectacular results can be achieved
with preparations that contain halogens.
It is very important to use the correct amount
of current for the encrusting anemones to do well
over the long haul. A medium to strong current
is advisable if numerous specimens are suffer-
ing badly from the deposition of detritus among
the polyps because of lack of current or too little
variation in the water movement. But a current
that is too strong and/or too direct can cause
the encrusting anemones to be stunted or fail to
open at all as a protective reaction. It can be as-
sumed that stunted polyps indicate too strong a
current, while very elongate, slender polyps re-
sult from lack of current. The same effect occurs
if the lighting is too strong (stunted polyps) or
there is a shortage of light (leggy polyps).

Only a very few zoanthids need to be fed deliber-
atelythe azooxanthellate species are one exam-
ple. Nevertheless, I recommend supplementary
feeding for the photosynthetic forms as well. In
many cases, growth and brilliance of coloration
can be further increased in this way. The trade
now offers a large selection of suitable ne pow-
dered or frozen foods, and it is ne to distribute
the particles in the water or add them into the
current. The direct feeding of individual polyps
isnt necessary and makes little sense unless you
are keeping very large Protoalythoa polyps or
propagating very delicate, slow-growing variants.
Zoanthids can be extremely toxic. This must always
be borne in mind when handling them. Avoid damag-
ing the polyps when taking cuttings, and always wear
gloves and protective goggles during such work. Despite
the thousands of import consignments that have passed
through my hands and the fact that contact with mu-
Left, top to bottom:
When these specimens from a tank dominated by blue
light were placed under a daylight fuorescent lamp,
they degenerated within a few weeks.
Snails of the family Architectonicidae, such as Psilaxis
oxytropis, may feed on encrusting anemones.
The Varigated Sundial Snail, Heliacus variegatus, is a
widespread predatory Indo-Pacifc species.
cous secretions from encrusting anemones hasnt always
been avoidable, I have never personally suffered poison-
ing of any kind. However, I am meticulous about wash-
ing my hands thoroughly after touching these animals.
Overall, I would say that the toxic effect of encrust-
ing anemones is appreciably more marked nowadays
than it was in earlier decades. This would also explain
the increase in problem incidents in recent years. We
can only speculate as to the causes of this, but I assume
that the abundant supply of high quality foods available
today may play a role. In the past, food preparations were
often lacking in essential fatty acids, and the products
available today might allow some specimens to thrive
so vigorously (even if they benet only indirectly from
the feeding of other corals) that they are able to gener-
ate their anti-predation toxins more strongly. However,
the fact that far fewer incidents of poisoning are known
from earlier decades may be due to the fact that many
cases of suddenly occurring, putative summer u were
actually the result of the inhala-
tion of a palytoxin-laced aerosol,
but werent linked to these poi-
sonous aquarium occupants. The
information most relevant for
the aquarist is that Zoanthus spe-
cies are less poisonous than their
relatives from the genera Palythoa
and Protopalythoa, which are usu-
ally responsible for serious poi-
There are various methods for the
propagation of encrusting anem-
ones. The procedure chosen de-
pends on whether the mother
is rmly attached to a substrate
rock or not. The most important
basic rule for long-term success-
ful propagation is to fragment
only healthy specimens that are
showing good growth. No matter
what the species and color variants, the cuttings should
be left in the original water for as long as possible, until
the almost inevitable damage from cutting has healed.
Fresh cuttings are extremely susceptible to bacterial in-
fections, so there should be no transfer or other change
in environment immediately after fragmentation. Buy-
ing fresh cuttings is not recommended.
The amount of time cuttings should be left in the tank
can often be gauged by the degree to which they have at-
tached to their substrate rocks. However, before plant-
ing, breeders should also take care to thoroughly clean
sponges and other biocover from the attachment surfaces
of substrate rocks that have already been in the aquari-
um, so that no decay takes place later beneath the join
or point of contact between polyps and hard substrate.
Cleanliness is very important in this type of work. For
your own safety, perform any fragmentation work involv-
ing zoanthids in a well-ventilated room and use protective
goggles, a breathing mask, and long gloves.
1. Fragmentation on rocks
For this method, a substrate rock with attached encrust-
ing anemones is divided into small fragments. The speci-
men should be rinsed in seawater thoroughly before cut-
ting in order to remove harmful detritus. It is important
that all the polyps are completely contracted, so that you
can target only the connecting basal tissue without cut-
ting the polyps themselves. Then, cut the low-growing,
shared body mass of the encrusting anemones through
with a scalpel, so that it doesnt tear when the rock is
divided, which can cause poorly healing wounds. A spe-
cial diamond band saw or diamond saw-blade (from
Dremel, for example) works best. With this method the
cutting is already rmly attached to a substrate rock, so
it is sufcient to attach it to a small substrate rock with
the aid of proprietary coral adhesive (as for stony cor-
als) such as one of the many super-glue gels. For pro-
fessional usage the most suitable frag mounts are those
with small pegs, with which they can be positioned in
cutting stands to grow on. The peg can easily be bro-
ken off prior to installation in the customers aquarium.
Good attachment and growing on after division can be
Aiptasia spp. anemones can sting and cause serious harm to
zoanthid colonies.
encouraged by the addition of a trace-element prepara-
tion containing a halogen complex.
2. Division of loose anemones
Ultra Zoanthus from Taiwan, Hong Kong, and the Ca-
ribbean are often imported without substrate. Once ac-
climated, they can be divided into cuttings. Because there
is no rock to cut through, you can use a scalpel. Make
sure it is very sharp and regularly replace the blade; sea-
water will quickly corrode the blade and make it blunt.
Loose cuttings must be attached to a cutting substrate
with a gel-type coral adhesive. This remains effective for
at most six months, but under optimal conditions the
zoanthid will have rmly attached itself by then.
3. Allowing encrusting anemones to spread
This is the easiest method of propagating zoanthids. A
small piece of hard substrate is simply placed immedi-
ately next to or even on or in a large mother specimen.
If the mother specimen is growing well, individual pol-
yps will soon start to grow on the small rock. Once this
happens the connection to the mother specimen can be
severed with a scalpel.
All in all, zoanthids have great potential for aquacultur-
ists. With systematic procedures and, for example, tar-
geted feeding, very large numbers of new polyps/cuttings
can be produced in a relatively short time. At Extreme
Corals (Germany) we currently produce
several hundred cuttings per month and
pass them on to the retail trade. The
high demand for them is undoubtedly
stimulated by the trend toward nano reef
aquariums, for which they are ideal occu-
pants. Unfortunately, not many profes-
sional coral breeders devote themselves
intensively to encrusting anemones, con-
centrating instead on SPS and LPS, but it
will surely be only a matter of time before
the aquarium trade recognizes the value
of these creatures on a broader front.
Like many corals, encrusting anemones
are susceptible to bacterial infections and
parasite attacks. I will give a short over-
view of the most commonly encountered
pathogens and parasites here.
Parasitic gastropods
(for example, Heliacus)
Various parasitic gastropod species
(snails and nudibranchs) are often
found on freshly imported stocks. They
can usually be removed fairly easily by
hand (this works with snails) or deci-
mated by the introduction of wrasses (for
example, Halichoeres cosmetus or Pseudo-
cheilinus hexataenia). Before introducing
new zoanthids into a reef aquarium, it is
advisable to keep them in quarantine for
a while so as to detect and remove any
parasites, or give them a preventative
bath in a suitable coral dip preparation.
The widespread and well-known Heliacus
snails can easily be recognized by their
shell form and conical operculum. Nu-
dibranchs, which are often adapted to
mimic the relevant encrusting anemone
perfectly, are appreciably harder to track
down. In such cases it is best to rely on the instincts of a
hard-working wrasse.
Zoa Spiders
These are sea spiders (pantopods), not true spiders as the
popular name might suggest. A few members of the huge
sea-spider classPycnogonida, with more than 1,300
speciesfeed on the tissue of encrusting anemones, and
they must always be found and removed prior to intro-
duction into the aquarium, as they may breed and cause
signicant damage. (They are not known to be venom-
ous and pose no threat to the aquarist.) In my opinion,
the safest method is a three-month quarantine with sev-
eral anti-parasite baths, which should take place prior to
selling or passing along new colonies.
White Fungus
This disease manifests via contracted pol-
yps that are coated in a white, foul-smell-
ing slime and eventually disintegrate. At
the rst signs of this problem, the af-
fected part of the encrusting anemone
should be cut away and removed (dont
forget to take safety precautions to avoid
contact with toxic slime!). Although the
popular name of the disease suggests
that a fungus is to blame, it is actually
a bacterial infection that usually results
from damage or stress caused by exces-
sively high temperature, salinity, or oxy-
gen shortage during transport. The chief
danger is that the infection will rapidly
spread over the entire colony if affected
areas arent removed immediately. In
many cases the bacterial attack may even
spread to previously healthy specimens
in the same aquarium. In the event of a
massive attack, treatment with Kanamy-
cin or Amoxicillin in an isolated hospi-
tal tank can be employed. In all cases,
existing areas of decay must be removed
and the affected polyp colony quaran-
tined, if possible in cool (7073F/21
23C), oxygen-rich water with weak
Although they are actually very easy to
keep, zoanthids quickly react to changes
in the aquarium environment. A change
in the brand of salt or slightly uctuating
water parameters can lead to long periods
during which the polyps remain closed.
This isnt always cause for concern; these
animals usually recover quickly from mi-
nor irritations. But if a zoanthid is driv-
en to keep its polyps closed because it is dealing with
constant upsets, a die-off might result. This can happen
if, for example, the polyps are repeatedly being stung
or nettled by Aiptasia or other actinians and cannot
escape their attacks. Fishes constantly nibbling at their
polyps will also cause this. In such cases the troublemak-
er must be removed without fail.
Excessively high temperatures or too rapid a reduc-
tion in food can cause encrusting anemones to bleach,
as stony corals do. Because they are relatively robust but
also rapidly exhibit external signs of being unwell, zo-
anthids are ideal canaries in the coal minethey will
often alert the aquarist in ample time to environmental
changes in his or her reef aquarium.
bsence makes the heart grow fonder, or so the saying goes. Of course that
doesnt apply in every case, and it most certainly didnt in this one. I rst
encountered the little gastropod called Aeolidiopsis harrietae, a member
of the family Aeolidiidae, in the spring of 1986, and it was my rst se-
riously negative experience in the reef aquarium hobby: they turned up on a newly
acquired colony of Zoanthus encrusting anemones and completely ruined them, as
well as another colony three months later. Eventually, the highly specialized mol-
lusks perished from lack of food.
I admit that I had considerably less enthusiasm for these mollusks back then
than I do today. At that time, I had recently entered the reef aquarium hobby and
was battling parasites with an uncontrollable reproductive rate that were eating my
actinians. I saw the gastropod as a parasite, a pest. Now, 26 years later, I see a mol-
Audacious gastropods
article and images by Dani el Knop
Sessile invertebrates
serve as a source of
food for numerous
creatures, including
encrusting anemones.
This article
demonstrates the
enormous effort a little
encrusting anemone
eating gastropod will
exert in terms of
mimetic adaptation to
avoid being discovered
by its predators.
Aeolidiopsis harrietae (right) doesnt hide,
but boldly curves its body into a rounded
form so that the cerata on its back
look like the tentacles of an encrusting
anemone. This fuorescent photo clearly
shows that green fuorescent pigments
are present in addition to the brown
coloration of embedded symbiotic
algae, just as in the host polyps. Below:
Encrusting anemones in the marine
aquarium, fuorescent photo.
lusk that has developed a fascinating mimetic
adaptation to its host actinian, and watch with
great interest as the gastropod curls its body
into a circle so that the appendages on its back
look like the tentacles of the neighboring en-
crusting anemones. Not only do they imitate
the brown coloration of the stolen symbiotic
algae that they accumulate, but they even pro-
duce a similar brilliant green uorescent effect
at the tentacle tipsa remarkable adaptation.
In the nal analysis, what makes these gastropods Trojan horses in the aquarium, where
they breed and harm others, is not wickedness but the aquarists ignorance. The mistake is to
put them in the aquarium with their host but without their own predators. This gastropod is
part of a highly complex system, a small cog in the overall mechanism, unable to turn prop-
erly by itself. Only if their proliferation is limited by predation pressure from their own preda-
tors will their effect on the host be limited and the system remain functional. To minimize
predation pressure on their own population, they have evolved their fascinating adaptation
to encrusting anemones. These days I am not battling against parasites, but watching a com-
munity of creatures that have evolved highly differentiated interactions and reciprocal rela-
tionships in order to survive, and I am fascinated by the ne balance they achievealthough
it doesnt always go smoothly in the aquarium.
I was pleased to be reunited with a once-hated enemy because I hadnt encountered Ae-
olidiopsis harrietae in 26 years, either in the aquarium trade or in the wild. It is known that
three members of the nudibranch family Aeolidiidae parasitize encrusting anemones, and all
three are in the genus Aeolidiopsis. A. harrietae isnt really uncommon in the wild, but it is so
well adapted to its host that it is rarely spotted. During the rst 10 years that I maintained
Left: Fluorescent photo
of an almost full-grown
specimen of Aeolidiopsis
harrietae; length is
.3 inch (8 mm).
Above: Two specimens
of Aeolidiopsis
harrietae of diferent
sizes suck on the same
polyp (normal and
fuorescent light).
Below: The sucking
process of the
gastropods leaves
behind a tissue
necrosis on the polyp,
visible as a raised
white spot.
coral reef aquariums, these gastropods were on my list of
aquarium enemies: they had robbed me of two beautiful
encrusting anemone colonies. In the second decade, by
contrast, I sometimes longed to have these gastropods as
alliesfor instance, when I experienced an uncontrol-
lable outbreak of the encrusting anemone Protopalythoa
heliodiscus in a 1,585-gallon (6,000-L) aquarium. But
despite a lot of effort and numerous communications
with dealers, wholesalers, and even exporters in Asia, no
specimens of this gastropod could be turned up.
During my third decade in the hobby I nally began
to see the interactions between the creatures in the coral
reef aquarium in a new way, with different priorities and
an eye for the interesting recip-
rocal relationships in an ecosys-
tem. This little gastropod had
become something of a teacher.
Aeolidiopsis harrietae belongs
to a group of nudibranchs that
have specialized on corals as food
and is fairly closely related to Ae-
olidiella stephanieae (Berghia),
which feeds on Glass Roses of the family Aiptasiidae. The
symptoms of a plague of gastropods are still the same:
the polyps of the encrusting anemone colony open in-
completely in a few places, and sometimes remain com-
pletely closed. Little whitish spots appear, sometimes
slightly raised and somewhat harder than the surround-
ing tissue. These are the places where the gastropods have
perforated the body of the encrusting anemone in order
to suck out its body uids. In the center of each spot is
a kind of plug composed of necrotic tissue. The gastro-
pods often hide among the polyps by day, but near the
closed mouth discs of the encrusting anemones you can
sometimes see the typical concentric clutches in which
the next generation of gastropods is developing. This spe-
cies develops without a planktonic stage, so the offspring
leave the eggs as fully formed gastropods, so tiny that that
The typically shaped clutches
of the Aeolidiopsis are found
on numerous polyps, and
the development of the
larvae can be seen with a
magnifying glass. By the
third day, microscopic
examination reveals
that the larvae possess
a concentrically wound
snail shell, but this has
disappeared by the time they
hatch on the sixth day. There
is no planktonic larval stage,
and the young gastropods
spread out on the encrusting
anemones and begin to eat
This photo of
the cerata under
magnifcation shows
the embedded brown
symbiotic algae and
the green fuorescent
they can barely be seen with the naked eye.
The manual removal of the gastropods and their
spawn can reduce the feeding pressure to some extent,
but may also damage the encrusting anemones. The
problem cannot be completely solved in this way, as the
young gastropods are innitely too small to spot them
without optical assistance. The larger ones can be col-
lected up daily, but new juvenile specimens will keep
turning up, seemingly out of nowhere, as they reach a
visible body size.
The aquarist is a less than perfect substitute predator.
The objective is far better achieved with the aid of natu-
ral predators that are specialized in tracking down the
young gastropods and have all day to do so. The encrust-
ing anemones suffer not from the presence of the gastro-
pods, but from the absence of their predators. It should
be easy to nd species that feed on these mollusks from
among the wrasses suitable for aquarium maintenance
(see box). It is, however, a prerequisite that the aquarium
also provide suitable habitat for these shes.
Knop, D. 2009. Trojaner im Meerwasseraquarium
Unerwnschte Aquariengste erkennen und bekmpfen. Natur
und Tier Verlag, Mnster, Germany.
Reef aquariums offer excellent conditions for nu-
merous small mollusks and annelid worms to re-
produce and spread, which isnt in the best interests
of the aquarist. In order to counter this, wrasses
should be added to the sh population, provided
that the environment satises their requirements.
It is imperative that they have an adequately deep
substrate in which they can bury themselves.
The choice of species is limited if you have a
smaller aquarium. The Six-Line Wrasse (Pseudo-
cheilinus hexataenia) is a possibility for volumes of
around 79 gallons (300 L) and up, though it can be
very territorial and aggressive at the adult stage. The
even smaller Pseudocheilinops ataenia is somewhat
more peaceful, but very rare in the trade.
There is a wider choice for aquariums that hold
132 gallons (500 L) or more: various Halichoeres
species can now be considered as well, for exam-
ple H. chrysus, which is relatively small and fairly
peaceful. H. cosmetus is signicantly less peaceful,
albeit also effective at eliminating small mollusks.
Even heavier predation pressure will be exercised by
the large-growing H. marginatus, but you will have
to expect a lot of disruptive behavior. Above all, the
larger the size of the wrasse in the adult stage, the
more they will deliberately overturn rocks in the
search for mollusks.
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Wheres the Beef ?
A nation of some
332 islands and
more than 500
islets, Fiji ofers
divers frst-hand
encounters with
many familiar
corals and reef
fshes seen in
marine aquariums.
Rainbow Reef and
Vanua Levu.
Right: Divemaster
Bale takes a shot
of Amphiprion
melanopus with
their Bubbletip
Sea Anemone.
7aveuni, August 2012 - 0ropping down to the reef, we pass through
a huge, owing school of blue and yellow fusiliers. I can feel and hear a con-
tinuous series of uctuating tones inside my head, and I check to see whose dive
computer is going ballistic. Other divers in the group seem equally puzzled, look-
ing this way and that. The divemaster gives a series of hand gestures I have never
encountered before, then takes out his regulator and, grinning, mouths the word
whales. It has been almost four years to the day since my last dive on Taveunis
Rainbow Reef, and Im being greeted by Humpbacks singing in the deep. It is not
hard to get excited about the diving in Fiji.
I rst came to Fiji in 2003 while backpacking around the South Pacic. This
small nation of islands, green and blue gems oating in an endless ocean, is one
of the friendliest places I have ever been. Generally safe and incredibly beautiful,
the country is full of fantastic people who will go out of their way to welcome you
to their island home. There are many interesting things to dodiving, sea kayak-
ing, surng, and sailing, as well as trekking and birdingand great cultural experi-
ences to be had. I have been many places, but few have called me back the way Fiji
has, and I have returned to her welcoming shores four times in the last nine years.
F i j
article and images by Nate Wi l son
Getting there is a bit of a hoof. Direct ights from
Los Angeles take about 10 hours. On the plus side,
they typically depart in the late evening and arrive in
Fiji in the early morning. If you are blessed with the
ability to sleep on planes, jet lag should not be a prob-
lem. Flights land in Nadi, on Fijis west coast.
We spent one day in Nadi organizing our travel to
Taveuni. To get from Nadi to Fijis capitol city, Suva,
my wife and I took the Coral Sun bus. We arranged
transport on a ferry from Suva to Taveuni the next af-
ternoon, an approximately 18-hour trip. (If time is a
concern, it is relatively easy to y from Nadi to the tiny
airport at Matei, on the northern tip of Taveuni.)
The air-conditioned Coral Sun coach picked us up
along the Wailoaloa road at 8:00 in the morning. We
followed the Queens highway south out of Nadi and
up into the mountains. Along the road the ame trees
were in blossom, crowning the green jungle with or-
ange ashes. Purple orchids grew along the roadside.
We passed through pastel-colored villages with taro
and bananas growing in the yards and saw beautiful
views of the coast and mountains. There was a brief
stop at a handicrafts market outside of Pacic Harbor,
and we were in Suva by 12:30. The bus stopped at the
Holiday Inn and we caught a cab down to the wharf
at Wali Bay, where we bought our ferry tickets. Our
boat, the Spirit of the Fiji Islands (SOFI), is an old Greek
car ferry that makes the trip to Taveuni, via Koro and
Savusavu, several times a week.
The ferry did not depart until afternoon, so we took
a cab back up to the National Museum of Fiji. There is a
small but interesting series of exhibits covering the his-
tory and culture of Fiji. The models of ocean-going ca-
noes are particularly fascinating. The feats of navigation
and sailing by open boat involved in the settling of the
Pacic were truly amazing, and the displays here call that
to mind. It is also evident that
the exhibits are put together
with a sense of national pride
and love.
On the SOFI, we had
sprung for a private room with
our own head, shower, and
queen-sized bed. There was
also an air-conditioned lounge
where videos played continuously. Absolutely knackered
from a combination of jet lag and walking around in the
equatorial sun, I promptly fell asleep. The night passed
quickly and we arose when the ship docked at Savusavu
around 6:00 A.M. We spent the next ve hours up on
the top deck enjoying the breeze as Vanua Levu, Fijis
second-largest island, slid by on the left.
Arriving by boat at an island, as visitors have done
for centuries, you get the chance to watch your desti-
nation slowly emerge out of the sea. Taveuni appeared
off the bow, rising from the Somosomo Strait, shrouded
in clouds and slowly growing longer and greener on the
horizon. Finally, after 39 hours of traveling, not includ-
ing layovers, we stepped onto the Government Wharf at
The Fish Factory, our rst dive site, delivers as advertised:
sh, sh, and more sh. To our left is the towering slope
of the reef, heads of hard coral mixed with patches of
the soft coral that abound here on the Rainbow Reef.
Currents in the Somosomo Strait between Taveuni and
Vanua Levu are cyclically strong and provide a constant
ow of nutrients. The underwater gar-
dens supported by such periodic currents
are astounding. We slide with the current
through shoals of anthias, hovering over
soft corals, zipping wrasses, and bright
parrotshes. Each tabling colony of hard
coral is home to schools of damsels and
Chromis; butteryshes or angels it along
beside us.
At our surface interval, we rest on
a white half-moon beach in one of the
shallow inlets of Vanua Levu. Taveuni lies
across the strait beneath a high, moun-
tainous spine draped in gray clouds. Our
boat rocks at anchor on gentle waves, in
turquoise shallows. The dive master and
boat captain pass out fresh bananas, tea,
coffee, and water, fried taro, and big, thin
Indian pancakes, called roti, spread with
honey and cinnamon. We all talk in a
rush, each member of the group eager to
talk about what he or she has just seen.
Everyone has something different to add:
the wealth of life on the reef is too great
for one person to catch all of it.
Diving the Rainbow Reef is like that. Each trip to the
bottom presents a mass of visual delight, impossible to
digest and process all at once. Swirls of small sh cover
the reef, and invertebrate jewels like nudibranches and
tiny shrimps are everywhere, awaiting closer inspection.
Look up and out in the blue are big pelagics hanging
out in the deeper water. Schools of barracudas, snappers,
and jacks drift in and out of sight along the reefs edge.
The occasional Dogtooth Tuna or Grey Reef Shark often
ventures up from the deep to inspect the reef. Napoleon
Wrasses and sea turtles drift across the reef tops.
After our snack break we head back out into the
strait. The water is like turquoise glass and the individual
coral heads can be seen clearly 60 feet below us. I am
itching to get in again. The divemaster, Bale, gives us a
quick brieng. He is a stocky Fijian with an infectious
grin and a mass of bushy hair with a peroxide blond
streak down the middle. When he isnt laughing, which
Above: Ring Wrasse,
Hologymnosus annulatus,
at Maxs Reef. This large,
famboyant species
is a threat to small
fshes and a wide range
of crustaceans and
A school of Golden
Sweepers, Parapriacanthus
ransonneti, swirls around
the reef at Nukubalavu
isnt often, he speaks in a rolling bass voice as if he is
channeling James Earl Jones.
Okay, this site is called Cabbage Patch, Bale says.
Some big green and yellow Scroll Coralit looks like a
cabbage. We are going to go down to about 50 feet [15
m], and you are going to see some hard corals and some
sh. We will swim over the Scroll Coral and then turn
left and follow the current down the reef. Keep the reef
on your right, and when you get to 100 bar, you let me
know. Then well start to go up and make a safety stop
for three minutes at 15 feet [5 m].
He neglects to mention that this mass of scrolling
coral is the size of a house and home to a bewildering
array of shestriggers, wrasses, parrotshes, squirrel-
shes, bannershes, and Chromis swirl over the gigan-
tic colony. After sufcient time to take in the massive
overlapping twists and scrolls, we n off through a maze
of coral pinnacles rising off the reef slope, and the dive
ends amidst mounds of fantastic soft-yellow leather cor-
als and an enormous boulder coral studded with hun-
dreds of brightly colored Christmas tree worms, their
feathery mouths ltering the currents for food. Silence
pervades on the boat ride back home. There is no more
of the rapid conversation of the surface interval. We are
all quiet, astonished by what we have just experienced.
I am diving with Jewel Bubble Divers, the only lo-
cally owned and staffed dive shop on the island. Qilolele
Morisio and his wife, Eunice, opened the shop in 2005
in a dive shack down the road from the present loca-
tion. My wife and I dove with them on our honeymoon
Anthias dance among the
sea fans and soft corals of
the Purple Corner.
in 2008 and the friendly, relaxed, and professional at-
mosphere created by the owners and guides, coupled
with the beautiful dive sites, made choosing Jewel Bubble
a no-brainer. I was pleasantly surprised to see that they
had moved into a larger space and added a second dive
boat and new rental gear. When Qio greeted us by name
as we walked into the shop, I was oored that he had
remembered us.
Jewel Bubble dives the Rainbow Reef, a 30- to 40-min-
ute boat ride down the Somosomo Strait, or closer sites
around Matei and Qamea. Over the course of two weeks
we went to the Rainbow Reef every day except one, when
the sea was choppy.
The next day we dive Nuku Reef, an impressive site
with mild current. My logbook entry reads sh and cor-
al, sh and coral, sh and coral. I come out
of the water eager to dive again and wonder-
ing what the Somosomo Strait will produce
next. The dive master laughs and promises
something special after the surface interval,
one of his favorites: Rainbows End. An hour
later I nd out why he is always smiling. We
pass through schools of fusiliers, which are
plentiful on every reef here, and into a can-
dyscape wonderland of layered hard and soft
corals stacked and piled together. We are sur-
rounded by Black Triggersh and Unicorn
Tangs, the ever-present orange icker of Sca-
len Anthias, and Blue-gold Fusiliersshes
that I will always identify with diving off
Taveuni. A lone Whitetip Reef Shark glides
away from us, and a Blue Ribbon Eel lurches
in and out of his burrow in the sand. The cur-
rent pushes us through shoals of parrotshes
and large tangs, over anemones loaded with
clownshes, above schools of bannershes
and Pyramid Butterysh, past zipping Xan-
thurus, Labropsis, and Jansens Wrasses and
a whole host of others that I cannot name.
In fact, the shes come and go too fast for
me to identify them all. Eventually I give up
and relax, letting the kaleidoscope of the reef
change and shift around me. The dive passes
quickly, and a little over an hour later I nd
myself waiting my turn to climb the ladder
back aboard.
The next day, Bale greets me with a hearty
Bula (the standard Fijiian greeting) as I
come down the driveway to the dive shop.
Last night it rained hard and the skies are
gray this morning. There is a steady wind blowing and
the seas are running choppy.
Bula, I respond. Looking out at the rough sea, I ask,
Rainbow Reef today?
No, no, Bale says, laughing. Today we go to Qa-
mea [Ga may uh].
Qamea is a large island just off Taveunis northern
tip. The ride is much shorter today, and bouncier. Devann
is the only other diver on the boat today; many canceled
because they thought the overcast skies presaged bad vis-
ibility underwater. Our persistence is rewarded by hard
coral forests and massive sea fans at Nukubalavu Point.
There is a heavy current running and big swells on
the surface, so we make a negative descent, dropping
as fast as safety will allow to the bottom. The reef here
hosts big red sea fans, and we land almost on top of a
particularly massive one with a huge school of Golden
Sweepers behind it. I spend a few minutes watching what
seems like thousands of these small sh ickering back
and forth beneath the outstretched scarlet lace. Bale
gets my attention by banging on the back of his tank
Top: Pink Anemonefsh pair with
their host anemone.
Above: Lemon Coral Goby rests
in Acropora branches near
Beverleys Beach, Taveuni.
with a metal pointer. Follow-
ing his outstretched arm, I see
Dogtooth Tunas racing through
the blue water and then realize
that a huge school of barracudas
hangs in the current, nearly mo-
tionless, just as interested in us
as we have become in them.
We surface-interval on Qamea, in a big horseshoe-
shaped cove. The beach is gorgeous, white, long, and
empty. Behind us, several houses are visible through the
palms and low brush. Devann and I swap dive stories
with Bale. He has been a dive master in Fiji for more
than 20 years; before Jewel Bubble he worked for a 7-star
resort north of Taveuni and then on a live-aboard dive
boat. He pays attention to us in the water, but you can
tell that he is still fascinated by the life on the reef.
A ve-minute boat ride takes us to Maraias Cove for
our next dive. It is shallow and loaded with
small shes. We discover a foot-long reef
scorpionsh while watching a pair of nu-
dibranches. There are clownshes nestled
in giant red anemones along with crowds
of single spotted black Dascyllus damsels.
Huge tabling Acropora share the reef with
soft, owery Dendronepthea. Chromis re-
treat into the hard coral heads as we ap-
proach. Beneath one I spot a huge purple
mantled clam. In another I am surprised to
nd a school of tiny yellow gobies with ma-
genta eyes darting amongst the Blue Green
The dive shop is closed on Saturdays, so my
wife and I decide to ride down to Nagara to
get fresh vegetables at the market. We catch
a bus just outside our accommodations.
The fare for the 30-minute trip is $1.70. Canvas aps
can be unrolled from the top of the bus over the open
windows to protect the passengers in the event of heavy
rain. It is a pleasant, breezy ride with postcard views. The
road hugs the coast, so on our jaunt down the island we
have the mountains and palm plantations on our left
and the rocky shores and blue water of the Somosomo
Strait on our right.
Saturday mornings in this friendly market town are
as close to hustle and bustle as you will get on Taveuni.
Farmers sell their goods from pickup trucks and wooden
stands along the main road. Taveuni is known through-
out Fiji as the Garden Isle, and on market mornings you
can see why. The stalls are awash in the greens of bok choi
and coriander, lettuce, and peppers, interspersed with
red tomatoes and hot chilies, orange squashes, yellow ba-
nanas, and purple eggplants. In 10 minutes we have ac-
quired enough vegetables to see us through the week. We
Above: Red-eyed lizardfsh
lies in ambush in the reef
Right: Reef garden with
pink soft corals, orange
sponge, and various table
Blackspotted Pufer
(Arothron nigropunctatus)
wedges itself between
pieces of the reef at low
tide of Beverleys Beach.
Coral Trout (Cephalopholis
miniata) nestles among
some of the red, purple,
and pink soft corals that
give Rainbow Reef its
retire to a small stall that sells tea, coffee, and breakfast
to wait for the bus. The woman behind the counter offers
us a delicious slice of pie lled with homemade pumpkin
jam. When quizzed about the recipe, she says that the
village women work together to make it. This mixture of
pineapple, pumpkin, and whatever else was handy when
it was made is a fantastic way to start the day.
My next dive is on one of Rainbow Reefs signature
sites: the Great White Wall. This site is so well known
that its listed in Lonely Planets guide to Fiji. It is the one
site on the Rainbow Reef that tends to be busy. Several
other boats are rafted up outside the reef when we arrive.
The good news is that their divers appear to be surfacing
as we make our giant strides into the water. This is the
only dive site where we have to share the water with an-
other boat, albeit briey.
To get to the wall you must n along the reef and
enter a swim-through at about 40 feet (12 m). The
tunnel is brief and you exit at around 70 feet (21 m).
I am pleasantly surprised to nd the current very mild,
and upon exiting the tunnel I am gently nudged along
through a cloud of big Golden Damsels and schools of
Square Blocked, Strawberry, and Scalen Anthias. Hover-
ing over a large Golden Sea Fan, Bale stops to point out
a pair of Longnose Hawksh perched within the jumble
of yellow branches. The wall is a massive vertical eld of
white soft corals pulsing in the currents. Aside from sea
whips poking out into the blue, there is nothing to stop
your descent, and paying attention to your depth gauge
is a must.
Toward the end of the dive we rise up and over the
wall through a small canyon in the reef. Somehow, Bale
manages to single out a small black pipesh hugging the
bottom amidst the chaos of corals and shes. Just be-
fore the safety stop we nd three lionsh hanging upside
down beneath an overhang crowded
with a school of jumbo Fine-Lined
The Great White Wall is a deep
dive, and we opt for a much shal-
lower dive for the second half of the
day. Jacks Place starts at around 50
feet (15 m), and we quickly climb the
reef to spend most of our time cruis-
ing along the reef top between 15 and
20 feet (4.56 m). It is essentially one
long and gorgeous safety stop among
stands of hard coral and swirling
masses of shes. This particular site
is one of the few on Rainbow Reef
where you can see blue Hippo Tangs.
Besides the smaller reef shes there
are also big schools of snappers, mul-
ticolored parrotshes, Unicorn Tangs,
and Clown Sweetlips cruising long
the reefs edge. The current is gentle
and allows an easy n among the ta-
bling coral gardens. At one bommie a
pair of big Sailn Tangs ashes away
among a shoal of Moorish Idols and a
lone Clown Surgeonsh. Titan Triggers
move around, digging at the coral with
their massive jaws. Anemones and at-
tendant Skunk Clowns dot the reef.
On a day with mild chop we hit
Purple Dreams and Annies Bommie,
both amazing dives.
I would take the end of Purple
Dreams over the renowned Great
White Wall any day. It is a dive that
sneaks up on you. Most of it is what
I have begun to consider average
Taveuni diving: a fantastic mixture
of hard and soft corals with the req-
uisite mass of small reef shes. The
dive plan carries us across several
reef slopes laden with coral and then
into channels of the reef with mini-
walls of soft white tree corals and
Purple Sea Fans. But these are not
the reason the site is called Purple
Dreams: after the last of the walls
the hard coral gives way to soft, and
the nal series of coral heads as you
near the surface are an explosion of
purple and lavender Dendronepthea
covered with a living veil of small
anthias. There is an enormous Gold-
en Sea Fan wedged into a mass of the
soft coral, and small orange Scalen
Anthias mix with slender Magenta
The frst of the three
waterfalls at Tavoro in
Bouma National Park. It
is a great place to swim
and there is enough room
to leap from the ledge
behind the cascade into
the deep pool below.
Anthias in huge shoals. As you near the purple corner,
the last protrusion before you pass onto the top of the
reef, the current strengthens, pushing you up and over or
around the point of pulsing purple soft corals. The pace
of the approach and sudden burst of color are absolutely
breathtakingand then it is gone.
I would have been happy to pack it in after Purple
Dreams, but Bale manages to follow it up in style. An-
nies Bommie does not disappoint. The dive starts with
a descent onto a at rubble eld studded with patches of
hard and soft corals. The gentle current pushes us down-
reef toward the rst bommie, rising to about 12 feet (3.6
m) below the surface. It is conical and wrapped in pur-
ple and red Dendronepthea. We circuit around the pin-
nacle through schools of anthias. Several small groups
of inquisitive goatshes and juvenile jacks swim down
out of the sun to check us out. Pairs of Regal Angelsh
and groups of yellow butteryshes move about the reef
without concern.
I would happily continue to circle this spot all day,
but now we bounce from side to side in a coral canyon be-
fore ending up on the right-hand side of the reef, passing
under an overhang where Red Coral Trout and an enor-
mous pair of morays look out at us. I spot three different
nudibranch species and we nish atop the reef, moving
through an enormous school of the ever-present fusiliers.
During a few days off, and we take an hour-long cab ride
down the other side of the island in a battered Toyota
minivan to the falls at Bouma National Park. The taxi
driver stops often so that we can get out and admire
views of the ocean or the jungle. The road turns to gravel
just north of Matei. I am surprised to be the rst one
signing the guest book for the day. A slight chill hangs in
the air at 9:00 in the morning, and we set off up the trail
with our swim suits, water bottle, and camera. We hike
the trails to all three of the Tavoro waterfalls.
The walk to the rst falls takes only 10 to 15 minutes
along a at trail. Here the falls spill over a cliff in the
jungle to a rocky pool 90 feet (27 m) below. You can
swim at all three falls and the water is crisp and refresh-
Huge schools of Magenta
and Scalefn Anthias
gather in the current to
feed above the soft coral
and sea fans at Purple
ing, especially after hiking. The second falls is another 45
minutes hike away. The initial climb is steep but not all
that long, and the local villagers are building a rest house
at the top of the hill. The trail splits off shortly there-
after; the left fork goes downhill and crosses a narrow
river before climbing up to the second falls. The right
fork will take you along the spine of the ridge through
jungle, all the way to the third falls. Unless there have
been heavy rains, the trails are fairly safe. They can be
muddy in places and the river crossing requires you to
hop across on rocks or get your feet wet. The views along
the trail, either back to the beach and Qamea or inland
to the mountainous center of Taveuni, are spectacular.
You will hear many birds even if you dont see them. We
spotted a small gray hawk hanging in the thermals at the
top of the rst climb, as well as a number of songbirds
and parrots throughout the hike. There are also purple
land crabs in the leaf litter of the forest oor and hun-
dreds of striped skinks with electric-blue tails. If nothing
else, walk up to the rst falls and have a swim.
On our way back from the third waterfall we stop to
talk with some men from the local village who are work-
ing on the rest hut. They offer us fresh coconut milk and
we share some of our cookies. There is a small visitors
center and canteen just off the Lavena road where the
bus stops across from the trailhead. It costs $15 to enter
the park. The men told us that in the old days, before this
was a national park, you paid $2 to someone in the vil-
lage and hiked up through the palm plantation. They tell
us there are visitors every day, but that it only gets very
busy during school vacations, when the local kids come
to the rst pool.
These men have hiked up the rst hill carrying sheets
of roong tin for the rest house. It has a commanding
view of the palm plantation and of the ocean, all the way
to Qamea. There are benches there and the checking out
the view is a good excuse to sit and recover from the climb.
I cannot imagine carrying everything for building a small
house up that trail one load at a time, but there is no other
way. The steps for the trails all the way to the second wa-
terfall are lled in with crushed coral pieces, which must
have been carried by hand, one bucket at a time.
Heading back, the trail is much easier than it was
coming up. The falls are visible through a circular hole in
the jungle canopy and invite us to push on. Down below,
we swim in the pool at the base of the rst waterfall. The
spray hangs silver curtains in the air and a circular cur-
rent sends small waves rolling into the rocks around the
pools edge. I climb the slippery rocks behind the falls
to leap off into the pool, and the cold water takes my
breath away. In the river, small shes that look like a
cross between carp and trout nibble at our feet. The wa-
ter is refreshing after our hike, and we swim for about an
hour. If you have made the effort to come all the way to
Taveuni and do not take the time to see Tavoro, you have
missed out on something truly fantastic.
LA57 0l V 0AY
On my last day of diving, the dive boat is full of fresh fac-
es. The dive master is kind enough to honor my request
to go back to the Purple Corner. It has rained throughout
the night and is just tailing off as we climb aboard the
boat for the ride to the reef. A low bank of clouds hangs
over the coast. Out over the water the sky is clear and
blue. A rainbow ends just off the point, plunging into the
turquoise waves of the Somosomo Strait.
Down on the reef the sky is gray and there are big
swells. We have to descend as soon as we enter the water.
Waiting on the bottom, we watch an octopus toss a crab
around like a basketball. The crab escapes several times,
but turns and menaces the octopus with his claws. His
eight-legged tormentor scoops him back up again and
again. Upon noticing us they both ee for shelter.
Visibility at the Purple Corner is even better today.
The tide is running in the opposite direction and the
corner seems to be in slightly deeper water, but is still
just as impressive. As we swim up the last wall, the soft
corals begin and even though I know what is coming, it
still takes my breath away: a mass of purple shrouded in
living sheets of shesclouds of Scalens and tiny Ma-
genta Anthias. The current is strong and whisks us up
and over. I signal to the dive master and swim back down
around the edge, peering into the mass of shes and col-
ors three more times.
Maxs Place is a great nish and a drift dive for sure.
I drop off the back of the boat into a mixed school of
Black Triggers, Unicorn Tangs, and parrotshes. The
strong current shoves me down the reef. I am literally
ying, stretching my arms out like wings, the reef pass-
ing below me at rapid speed. The current carries our
group over the reef through a little cut and then tapers
off as we hit the other side. There are Blue Ribbon Eels
on the bottom and schools of fusiliers with electric blue
spots, Sweetlips, and Snappers moving ahead of us, along
with squirrelshes, a pair of giant bannershes, and a
Napoleon Wrasse. For about ve minutes, a Green Sea
Turtle swims among our group, pacing us over the coral.
As I pass up and off the reef a lone parrotsh, eeing
the other divers in our group, poops a blast of chewed-up
sand on top of me. It is if the reef is saying goodbye. Then
the blue water, a safety stop amid fusiliers, the drift line,
timing the swell to get back on the ladder, and a slow
30-minute boat ride back up the strait, running toward
Matei and Taveuni, shrouded in clouds. I am quiet, re-
ecting on the dives of the past two weeks.
That night we enjoy a feast at the home of our host
family. We share kava and stories, fried sh, chicken cur-
ry, cassava and rice, my wifes Alloo Ghobi (spiced pota-
toes and cauliower), and an eggplant stir-fry. Walking
back to the bure under the Southern Cross, I can see,
through the avenues of palm trees, the sea inging the
diamond-bright stars back into the Milky Way. No won-
der people are drawn back to Taveuni.
Unnoticed by many marine aquarists,
copepods are astonishingly complex
crustaceans with anatomies built for
predator evasion. Color-enhanced macro
image by Igor Siwanowicz.
Up Close
Monospecic aggregations of copepods were ob-
served and sampled from 11 coral reefsin the
Central and Northern Great Barrier Reef Prov-
inces of Australia and in Palau, Western Caro-
line Islands. Acartia australis, Acartia bispinosa,
Oithona oculata, and Centropages orsinii regularly
occurred at swarm densities of from 500,000
per cubic meterto 1,500,000 per cubic meter
Swarms of 3570 cubic feet (12 cubic m) were
common; they occasionally exceeded 1,060 cu-
bic feet (30 cubic m). Continuous linear swarms
of 65.5164 feet (2050 m) were common. We
have seen swarms longer than 328 feet (100
m), giving an upper limit on total swarm vol-
ume of about 1,766 cubic feet (50 cubic m). At
1,500,000 O. oculata per cubic meter, the total
number of copepods in such a swarm would be
75 million.
by Ronal d L. Shi mek, Ph. D.

Hamner, W.M. and J.H. Carleton. 1979. Copepod swarms:
attributes and role in coral reef ecosystems. Limnology and
Oceanography 24: 114.
As I hope will become apparent, very good arguments
may be made for considering an average copepod to be
the average earthling of the animal persuasion. Virtually
all aspects of their natural history are remarkable, and
simply because of their immense numbers, even what of-
ten appear to minor things turn out to be not so minor.
Finally, they are notable for having their major attributes
and properties continually studied and totally miscon-
strued or misinterpreted by some of the best scientic
minds available.
As just one example of the inconsequential being
anything but inconsequential, the pelagic copepods di-
urnal behavioral pattern of ascending from deeper water
to the shallows and subsequently returning to the dim,
dark depths, covering at most a vertical distance of a few
thousand feet, seems relatively trivial. However, with a
little thought it becomes evident that it is anything but
triing. When multiplied by the number of animals that
make it, that little round trip has rightly been described
as the largest migration known; it moves hundreds of
billions of animals over a regular migratory route on a
daily basis. While the distance doesnt seem like much to
us, it is a long distance to a small crustacean; worldwide, T


of Acartia
sp. calanoid
an order that
dominates the
numbers of
in the worlds
the amount of biomass moved is absolutely staggering
and the periodicity is precise. At other times, the move-
ments of a single tiny animal may be just as remarkable;
individuals of the calanoid Rhincalanus nasutus move at
speeds in excess of 400 body lengths per second during
their escape responses. A human moving with the same
proportional velocity could easily outrun bullets.
Most reef aquarium hobbyists probably know what
copepods are; they are those little white dots that move
slowly in the water, or with jerky motions across the
aquarium wall. UhhuhWell, yeah. And then some. I
hope this article will serve as an introduction to the basic
pod of all pods, the critter that is the exemplar of all
small crustaceans. Some observers credit copepods with
being the largest animal biomass on earth, and, as with
krill, they are considered one of the key carbon sinks in
the ocean, consuming phytoplankton near the surface
and depositing their wastes and molted exoskeletons on
the bottom at depth.
A noted scientist who lived around the middle of the
last century, J.B.S. Haldane, is said to have given a pre-
sentation to a group of seminarians, after which he was
asked, What can you discern about the Creator from
your study of creation? Haldane, an atheist, is said to
Largest to Smallest: Parasitic copepod forms tend to be larger than free-living
ones, and the largest copepods are Pennella individuals about a foot (30 cm) in
length that live with their heads embedded in, and feeding on, whale blubber.
The rest of the copepod extends out of the whales skin and looks like a long,
thick, brown string. The largest free-living copepod known is a .71-inch (18
mm) female Bathycalanus svedrupi, a deep-water calanoid species. The smallest
adult copepods are unknown, but are probably free-living. Our ability to sample
very small live animals is rudimentary, and most of the oceans smaller animals
are probably still unknown.
Largest Migration: Individuals of Oithona species, oceanic cyclopoid copepods
with an adult size around 0.1 mm, swim from 1,650 feet (500 m) to the surface
and back down again daily. Their numbers are incalculable; up to 70 percent of
all plankton from 60N to 60S latitude move in this manner, and most of that
plankton is comprised of Oithona individuals.
Largest Number of Individual Animals in a Swarm: Swarms are animal groups
that move with synchronized motion, such as bird ocks and sh schools. The
largest measured copepod swarm occupied over 1,059 cubic feet (30 m
) and
was estimated to contain over 30,000,000 animals, all swimming in a synchro-
nized manner.
The Strongest Anythings: Calanoid copepods have been shown to be more than
1030 times stronger than any other animal in force production per unit time,
due to the rapidity and sheer strength of their muscle contraction. Per unit
weight, they are also stronger than any man-made device.
The Fastest Animals Relative to Their Size: A 1-mm copepod moving by a muscle
twitch moves at the rate of about 20 inches (.5 m) per second. If that motion
were scaled up, a human swimmer in what, to a copepod, is thick, gooey, viscous
water would spurt about half a mile (805 m) in a second, or a velocity of around
1,800 mph from one kick, which unfortunately would atten the unprotected
swimmer to microscopic thickness. If the human jumped in air rather than wa-
ter, and accounting for differences in the two media, primarily the 1,000-fold
difference between their densities (water 1,000kg/m
; air 1.3kg/m
at sea
level), a human with the proportional strength of a copepod jumping up would,
quite simply, not come down. The person would reach a velocity between 6 miles
(9.6 km) per second (that necessary to achieve high earth orbit) and 12 miles
(19 km) per second (well exceeding solar system escape velocity).
Birtles, A., and P. Arnold. 2002. Dwarf
minke whales in the Great Barrier Reef
CRC Reef Research Centre, Current State
of Knowledge, May 2002.
Gallienne. C. P. and D. B. Robins. 2001.
Is Oithona the most important copepod
in the worlds oceans? J Plankton
Res 23 (12): 142132. doi: 10.1093/
B. Gemmell, H. Jiang, J.R. Strickler,
and E.J. Buskey. 2012. Plankton
reach new heights in efort to avoid
predators. Proc Roy Soc B, http://dx.doi.
Hiskey, Daven. 2010. http://www.
John Mauchline.1998. Introduction,
in The Biology of Calanoid
Copepods. Vol. 33, Advances in
Marine Biology, pp. 115. Elsevier.
Kirboe, T., A. Andersen, V.J. Langlois,
and H.H. Jakobsen. 2010. Unsteady
motion: escape jumps in planktonic
copepods, their kinematics and
energetics. J Roy Soc Interface. doi:
Buskey, E.J., J.O. Peterson, and J.W.
Ambler. 1996. The swarming behavior
of the copepod Dioithona oculata: In
situ and laboratory studies. Limnol
Oceanogr 41 (3): 51321. doi: 10.4319/
Copepod Extremes







have responded, If one could conclude anything about
the nature of the Creator from a study of creation, it
would appear that God has an inordinate fondness for
stars and beetles. Haldane, like any good public speaker,
knew a good phrase when he uttered one, and he ap-
parently used the phrase, an inordinate fondness for
beetles in several different ven-
ues, including, at least once, in
print. Later, reecting on the large
number of described beetle species
(about 400,000 in Haldanes day,
around 750,000 in 2010), he is
reputed to have added, What was
it about beetles that took Him so
many tries to get them right? After
all, God could get man right on the
rst try, or at least we presume so.
There is not a shred of a doubt
that the insect taxonomic group re-
ferred to as the Order Coleoptera,
the beetles, takes the prize for hav-
ing the largest number of species. But while there may be
more species of beetles than of any other type of animal,
beetles are absolutely, and without a doubt, not the most
numerous type of animal; that distinction belongs to
the copepods. By some estimates, there may be as many
as 15,000 copepod species, a rather piddling number
compared to the beetlesthere are roughly 50 species of
beetle for each copepod species. Still, copepod individu-
als outnumber the beetles, probably by an unimaginably
large margin.
How was that useless piece of information calcu-
lated? Well, rst, one persons useless is another per-
sons interesting. Given that the oceans are fairly poorly
known, it is a given that they have not been adequate-
ly sampled for any group of organisms, let alone spe-
cically copepods. Nonetheless, a noted copepodiatrist,
G.A. Boxshall, made what could be called a reasonable
ballpark estimate, or a WAG (Wild A = Small Donkey
Guess). First, noting what makes our pale blue dot blue,
Boxshall said that 71 percent of Earths surface is cov-
ered by a thin water layer averaging about 12,100 feet
(3.7 km) deep, creating a volume of 1.347 billion cubic
kilometers. A single cubic kilometer contains 1 trillion
(1.0 x 1012) liters, meaning that the oceans contain
precisely 1.347 x 1021 liters of water. Assuming that
the average oceanic copepod abundance is only one
individual per liter, which is probably a signicant un-
derestimate, there would be 1,347,000,000,000,000
copepods swimming in the seas. Not only is this num-
ber unimaginable, it is unimaginably larger than any
estimates of the number of beetles or, for that matter,
of all insects combined.
However, Boxshall did his back-of-the-envelope
calculations in the mid-twentieth century, and it has
since been determined that the number of copepods
were drastically underestimated due to improper sam-
pling, mostly in the tropics, specically around coral
reefs (see the Hamner and Carleton article quoted at the
beginning of this article). Starting in the 1970s, (mostly)
young researchers using scuba gear, as well as other dif-
ferent sampling methodologies, decided to break with al-
most a century of tradition and get in the water and ac-
tually observe (GASP!!) and document what proportion
of the plankton was sampled by plankton nets around
coral reefs and elsewhere, as opposed to just sampling
the waters and blithely assuming the samples were ad-
equate and unbiased.
These studies resulted in a wholesale revision in
sampling techniques. Using the new methodologies,
the number of copepods typically recorded in samples
jumped by a factor of 10 to 100 times. Over a coral reef,
where previously a researcher would expect to nd from
300 to 3,000 copepods per cubic meter of water, more
recent samples found upwards of 300,000 individuals.
As if recording only one tenth to one hundredth of the
numerical total of the planktonic animals wasnt bad
enough, many species of small copepods were com-
pletely missed in the older samples. Having adult sizes
of 100 m or smaller in many cases, these species actu-
ally turned out to be the numerically dominant constitu-
ents of the plankton. Interestingly, examination of some
older preserved, but uncounted samples collected in a
different manner for different purposes showed without
Harpacticoid juvenile
copepod resting atop an
Artemia cyst.
Harpacticoid copepod is minuscule but displays
the exoskeleton and bristling anatomy typical
of many crustaceans.
Pleuromamma, a calanoid
copepod captured by
a scanning electron
microscope with colors
applied to illustrate
diferent body parts. Note
the hair-like setae on the
long antennae, vital in
detecting odors and in
managing motion.


Gaussia, at 6
mm a giant
among marine


a doubt that the missing critters were present when the
previous samples were taken. They were simply missed
because the samples were taken in the wrong manner.
The bottom line is that the worlds marine copepod pop-
ulation can now be conservatively estimated to be 1.35
million trillion animals. Counting them at one animal
per second, it would take 42,713,089,802 years, 47 days,
18 hours, 41 minutes, and 2.4 seconds, or roughly 3.1
times the time theoretically considered to have occurred
since the Big Bang started the universe on its way, to
count them alland thats not including time for lunch-
es or potty breaks.
The shift in copepod numbers resulting from the re-
evaluation of sampling methods has resulted in another
interesting change in the les of useless trivia. On the
basis of their numbers in the plankton of the bo-
real and temperate seas, large (24-mm) copepods
in the genus Calanus had long been thought to be
the most abundant animals in the world. Now, on
the basis of their numbers in all seas, the smaller
(adult length 0.1 mm) cyclopoid copepods in the
genus Oithona are now considered to be the most
numerically abundant animals. I suppose that I
might be accused of splitting hairs, but really, con-
sidering the group, it is copepod setae (bristles)
that are being split. And given that Oithona and
many other copepods are often very plentiful over
coral reefs, the concept that coral reefs were liv-
ing in plankton-poor seas has rightly found its way
into the trash bin.
Derived from the Greek (= kope: handle or
oar) and o (= pod; foot), copepod means oar
foot and refers to the swimming motions made by
the pairs of thoracic appendages. Each pair of append-
ages works together, like a pair of oars, to provide steady
and consistent locomotion.
The Copepoda, having about 12,000 described spe-
cies, are the second most species-rich crustacean group
after the Malacostraca. Five of the ten taxonomic or-
ders, or major subdivisions, of the Copepoda contain
species that may be entirely free-living. Most free-living
copepods are found in the orders Calanoida, Cyclop-
oida, and Harpacticoida, with only a few species found
in each of the other two orders, the Gelyelloida and the
Platycopioida. Animals from the latter two orders, and
the species contained in the ve additional taxonomic
orders having only parasitic forms, will be ignored in the
rest of this article.
Dorsal view of a
copepod showing
muscles, gut, and
eye (a light-
sensing organ).
Life-size, many
marine copepods
in the aquarium
are no larger than
motes of dust.
Female copepod with developing
mass of nauplii (arrow).
Unfortunately for a survey article such as this, the cal-
anoids, cyclopoids, and harpacticoids are neither closely
related nor particularly similar in their morphology and
natural history, and the research about them is scattered.
The calanoids, the focus of most copepod research, can
be generalized as being marine and pelagic. Cyclopoid
research has been concentrated on freshwater species,
although the cyclopoids are also mostly pelagic and very
common in marine systems. With the recent discovery
of the awesome numbers of small under- and unsampled
marine cyclopoids in genera such as Oithona, that effort
will undoubtedly change. Compared to the cyclopoids
and calanoids, harpacticoids are not often studied, pos-
sibly because they have a reputation for being difcult to
identifybut most appear to be marine infaunal or epi-
faunal benthic crawlers or near-bottom swimmers. There
are also a number of parasitic forms found in this group.
The three major primarily free-living copepod types differ
in appendage shapes and relative prominence and where
the abdomen articulates with the rest of the body.
or with one, unfortunately. Although they are often
quite attractive and have exceptionally interesting attri-
butes and natural history, their tiny size, coupled with a
plethora of appendages, can make copepods essentially
impossible for a non-specialist to even understand, let
alone deal with. However, for the person willing to invest
some time reading and to spend the time and money to
build or obtain the means to observe them, these micro-
scopically mesmerizing crustaceans can truly open the
door to those unseen parts of the coral reef that most
aquarists never seethe plankton and the meiofauna
(organisms that can pass through a 1-mm mesh). For
the aquarist who knows the reef, the copepods become
the raison dtre of the natural reef; in short, they feed
corals as well as many shes and other invertebrates.
Without them, the reef would die. For that reason, let
alone their own intrinsic and interesting attributes, an
aquarist should nd them worth learning about.
The standard-issue copepod body is a tapered or tear-
drop-shaped cylinder containing 17, or fewer, segments,
although some segments are not clearly delineated. Co-
pepods lack heads, but the ve anterior-most pairs of
appendages, often referred to as head appendages, are
all present; these, and the appendages from one or two
more thoracic segments, are fused to form the anterior
body region, the cephalosome. The ve appendage-bearing
segments of the thorax comprise the middle of the ani-
mal, and bringing up the rear is the abdomen, which has
up to ve segments, all lacking appendages, but ending
in a segment with two branches, the caudal furca. The
thoracic segments are often referred to as the metasome,
and the abdominal segments form the urosome. In some
copepods, more amalgamation between the cephalo-
some and metasome results in a larger, more inclusive
body region called a prosome. Fortunately for a person
wanting to learn about them, free-living copepods are
bilaterally symmetricalthe animals paired appendages
are mirror images of one another. Free-living adult co-
pepods vary in size over about two orders of magnitude,
from about 0.1 mm to about 20 mm. Some of
the parasites may be a bit over a foot in length.
It is sometimes useful to think of crusta-
ceans as the Swiss Army knives of the ani-
mal kingdom. Within each species, each pair
of appendages is different from all the other
pairs, and what make each species unique are
the modications seen in each limb pair. The copepods
buck this trend to a small extent, but there is enough ap-
pendage diversity within and between the various groups
to drive a person buggy. As with all crustaceans, at least
a partial key to understanding the whole animal is know-
ing the functions of each appendage.

Misconceptions about copepod natural history got their
start right at the beginning of ocean science. Large cala-
noid copepods were some of the rst marine animals to
be discovered at the birth of oceanography in the late
nineteenth century. Particularly common in the north-
ern seas where oceanography was developed by pioneer-
ing Danish and German scientists, these animals were
obvious targets of study. It didnt hurt that many of them
were also quite beautiful. Common, pretty, and cheap
what better animals to study?
Examination of the functional morphology of the
appendages of the large calanoid copepods shows that
the thoracic appendages and those around the mouth,
the so-called mouthparts, could only function in one
way. One look at all those appendages with their lobes
fringed in long, stiff setae, and it was instantly obvious
the animals used these appendages to sieve the water for
phytoplankton, much as a North Sea sherman uses a
net to harvest herring. It really was trivially easy to de-
duce how copepods fed. Anybody could visualize himself
as a copepod, swimming in a pool, using something like
a short, hand-held leaf rake to simulate the setae on his
appendages. If some neutrally buoyant rubber balls rep-
resenting the algal cells were added, the mechanics of
copepod feeding were intuitively obvious: they dragged
the setae through the water to lter out useable food
items. Thirty years ago, or thereabouts, that is how this
section of this article would have ended. As with sam-
It is sometimes useful to think of crustaceans as
the Swiss Army knives of the animal kingdom.
pling of plankton, nobody looked in detail at just how these animals actually
were feeding.
It turns out that many pelagic copepods feed in a manner wholly unlike any-
thing anyone expected. From the beginning of ocean science and for the next
century or so, it remained such a certainty that the calanoids caught their
food by lter-feeding that when Mimi Koehl and Rudi Strickler published
their 1981 paper detailing exactly how calanoid copepods actually fed, the cu-
mulative surprise evinced by oceanographers and copepodophiles everywhere
was truly watershaking.
Using a deductive mental experiment, such as that initially used to ex-
plain copepod lter-feeding, often allows a quick jump to an accurate con-
clusion. Unfortunately, if some of the many intuitively obvious factors are
neither truly intuitive nor inherently obvious, that quick jump can be more
like the lemmings proverbial quick trip off the edge of a cliff. In this case, the
thought experiment of humans cavorting in a swimming pool turns out to be
quite unlike the actions of copepods swimming in the ocean. Because of their
small size and low mass, copepods experience a water world vastly different
from anything humans experience.
Inertia and viscosity, properties of items immersed in a uid and the uid
itself, respectively, are determined by the size and the mass of the object.
Humans are massive animals, which gives us a great deal of inertia. When
a human swimmer quits ailing at the water, she typically coasts for some
distance before she stops. If a copepod stops its propulsive motions, it stops
immediately. It doesnt coast, because it cant coast; it is so very light that
it is effectively massless and has almost no inertia. Besides, the water ows
past a human swimmer. There is some friction with the water, enough so
that competitive swimmers wear specially designed swimsuits that allow the
water to slide past them with a minimum of drag. The water doesnt slip past
a copepod; it sticks to it. To experience what it would be like to swim in a
copepods world, a human would have to be swimming in a uid with the
viscosity of cool molasses or honey.
In the world of small sizes, where things effectively have no inertia and where
viscosity rules, trying to snag a meal can be a real problem. Because of the
viscosity, water will not ow between the bristles on the tip of an append-
age, so sieving is just not possible. A calanoid uses its second antennae, part
of its mandibles, both pairs of maxillae, and the maxillipeds to feed. When
the animal feeds, the second antennae and the maxillipeds simultaneously
ex outward, toward the front and rear of the copepod, respectively, pushing
some water to the front and rear and simultaneously pulling a blob of water
containing some algal cells inward, between them, toward the body at the
level of the maxillae. A sequence of maxillary movements shoves much of
the incoming water blob laterally and posteriorly, while corraling a portion
of the water mass containing the algal cells. Water is deected, scraped, or
pulled away from the food/water mass by the maxillipeds and both pairs of
maxillae, and the progressively smaller algal-containing water blob is moved
forward toward and into the mouth. By the time the algal cells are actually at
the level of the mouth, they are surrounded by a very thin lm of water, and
they often have not been physically touched by any part of any appendage.
No water has been ltered through any mesh; the water has been sliced
off, pushed aside, or allowed to ow between appendages. The appendage
lobes behave more like paddles, or spatulas, than like rakes or strainers during
this process. In short, copepods do not lter-feed; they use their phenomenal
sensory equipment to detect and isolate prey items with-
in a nearby water mass, then actively isolate those parcels
of algae-containing water, progressively shaving the
water away, leaving only the algal cells to be eaten.
Variations in the appendage movements can result
in unacceptable particles being rejected, and differences
in appendage shapes result in different prey selectivity
by different copepod species or growth stages. Of course,
not all pelagic copepods are phytoplankton predators;
some prey on living animals. In the pelagic copepods
world, things are small, but they are not placid or se-
date. Relative to their size, predatory copepods are prob-
ably more fearsome than lions or tigers. They are rapidly
moving predators that subdue prey not much smaller
than themselves, and in addition to having strong rap-
toral appendages, at least some of them are venomous.
The marine habitat contains a lot of copepod species,
so it follows that those that live most closely together are
likely to have different diets or feeding habits. Benthic or
sediment-dwelling harpacticoids are generally presumed
to glean food from the bottom, either scraping it from
rocks or sand grains or harvesting tasty bits of decom-
posing something-or-other. How they actually go about
this is distinctly unclear at present. Given how inaccu-
rate the model of lter-feeding was, it would be foolish
to assume we know how the gleaners/grazers feed.
Many types of animals have a non-living outer layer
completely covering the animals body. Often called a
cuticle, this layer is secreted by the often thin, outermost
tissue layer, the epidermis. Cuticles may sometimes be
elaborated into structures such as the familiar fur, feath-
ers, or teeth of vertebrates. In echinoderms, this cuticle
is so thin that it often takes a transmission electron
microscope to generate the magnication necessary to
demonstrate it. In the case of arthropods, however, the
cuticle really gives the animal its form. In animals such
as the copepods we are considering here, the cuticle and
some other underlying structures form the exoskeleton,
or better, the integument, which, in addition to providing
a system of levers to allow efcient muscular movement,
literally provides a form for the animal. Without the in-
tegument, there would be no crustacean.
The arthropod integument/exoskeleton is truly a
marvel of bioengineering; it is exible, yet impermeable
to chemicals other than water and dissolved gases. When
thin and in its basic form, it is absolutely transparent,
but the addition of pigments opens up a world of colors.
Copepods are often considered to be white or colorless,
but oceanic copepods run the gamut from cherry red to
cobalt blue.
The cuticle doesnt stretchat all! For the animal
to increase in volume, the integument must be periodi-
cally shed and a new one secreted. And one of the several
things that can make this molting process REAL
LY dicey for the bug is that the tissues lining the foregut
(the esophagus and the anterior part of the stomach) and
the hindgut (what passes for the rectum) are lined with a
tissue layer that is modied external skin, and are, like all
external surfaces, covered with an exoskeleton layer. So
for the animal to grow and molt, it must not only shed
its skin, it must also detach the thin exoskeleton lining
of the esophagus, stomach, and hindgut and pull those
linings out through the mouth and anus, respectively. All
of this must be done without tearing the delicate sur-
rounding tissues that are now deprived of their support-
ing exoskeleton. The process must occur without the lin-
ing getting hung up on some of the other guts,
gadgets, or appendages comprising the animal.
As might be imagined, this process is not
undertaken lightly. Judging by the crustaceans
whose growth has been examined closely, these
animals probability of dying during the molt-
ing process varies from molt to molt, ranging
from less than 1 percent to upwards of several percent.
Molting is controlled by the copepods endocrine system,
and in those species that live long enough, variations
in day length also exert some control. As with all other
crustaceans, iodine has nothing to do with molting.
Crustacean integuments have a very complex com-
position, but the major components are layers of sev-
eral durable proteins, lipids (possibly for waterproong),
and a rather strange sugar polymer called chitin, which
is responsible for the exoskeletons remarkable proper-
ties. Chitin is similar to the well-known structural sugar
polymer cellulose, but differs in that its sugar subunits all
have an attached ammonium residue. Chitin is almost
chemically indestructible. In the formative days of mod-
ern biology, in the latter half of the nineteenth century,
it was easy, but hazardous, to test for chitin. The sample
thought to contain chitin was boiled in concentrated
sulfuric acid for 24 hours. The uid was poured off and
any residue was rinsed and washed with distilled water.
Then the sample was boiled for 24 hours in a maximally
concentrated lye solution. Anything remaining after this
treatment was chitin.
Today, there are less hazardous ways of testing for
this substance, but no matter what method was used, it
would be found in copepod skeletons. Larger crustaceans
often have a waxy outer layer and a calcium carbonate
inner component, but both are generally lacking in co-
pepods. All integumental components are secreted by the
epidermal tissue, sometimes referred to as the hypoder-
mis (hypo = under), that lies under the non-living layers
The arthropod integument/exoskeleton is truly
a marvel of bioengineering....
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of the integument.
Solving the problem of pulling the shed integument
out of the ends of the digestive tube should be simple.
Since the copepod secreted the chitin in the rst place, it
should be able to dissolve it and simply slough off the tis-
sues. However, virtually no arthropods have either cellu-
lases or chitinases, the enzymes that dissolve cellulose or
chitin, respectively. Most wood-eating arthropods, such
as termites, have bacterial symbionts that digest cellu-
lose, but no arthropod has ever been demonstrated to be
capable of digesting chitin.
Although many copepods, particularly the small-
er harpacticoid copepods, are the right size for ciliary
propulsion, the cuticular covering of all external body
surfaces rules this out. No arthropod has motile cilia,
although immobile cilia, found mostly in sensory struc-
tures, are common. Nevertheless, all copepod locomo-
tion is done by muscle action, which raises one more
integumentary problem: muscles transmit their forces by
being attached to various skeletal components that func-
tion as levers. During molting, the muscles must be de-
tached from the old skeleton so that it may be discarded,
then reattached to the new skeleton so that the animal
can move. Consequently, there are periods during molt-
ing when the newly molted animals exoskeleton has not
yet hardened, so it cannot move well, if at all.
In large crustaceans, such as crabs or shrimps, it
takes one or more post-molting days for the integu-
ment to chemically cure and become sufciently rigid
for muscle action to move the animal instead of sim-
ply deforming the exoskeleton. Pelagic copepods dont
have the luxury of time; without the ability to move,
they cant avoid predators. In most copepods the newly
secreted skeleton appears to harden very rapidly, often
within a minute or two. The physiology of this remark-
able process is by no means understood.
Pelagic oceanic copepods are the epitome of small, high-
ly muscular animals that move exceptionally rapidly for
their size. The harpacticoids that live on and between
sand grains are by no means less muscular, but they are
adapted for a different type of motion. Yet both possess
the same type of musculature, on the microscopic level.
Copepods are capable of moving with precise move-
ments, whether it is with slow, delicate deliberation or
with high-speed reaction. Such motion is the result of a
degree of neuromuscular coordination that has seldom
been documented in other invertebrates. Presumably,
the larger pelagic copepods live in a system where preda-
tion is always imminent, and escape must be extremely
rapid. This has resulted in natural selection working to
allow exceptionally fast reactions and responses, particu-
larly for such small organisms.
And how do copepods respond to predators? Plank-
tonic ones try to get rapidly out of the way. Rhincalanus
nasutus from the Indian Ocean, a relatively large cala-
noid, about 5 mm maximum length, has been measured
moving at an amazing 6.8 feet (2,085 mm, or about 420
body lengths) per second! A 6-foot (1.83 m) human
moving at the same relative speed would have a velocity of
about 2,500 feet (762 m) per second, which is faster than
the muzzle velocity of the bullets of virtually all handguns
and many ries. Some other calanoid species, in a group
called the pontellids, normally live in the upper few milli-
meters of water. Two Gulf of Mexico species, Anomalocera
ornata and Labidocera aestiva, each about 3 mm in length,
are preyed upon by small shes called mullets. Individual
copepods, sensing the pressure wave generated by an ap-
proaching sh, generate a powerful swimming
stroke by rapidly contracting all of their thorac-
ic legs in succession, simultaneously stream-
lining themselves by pulling both antennal
pairs tightly to the body; as a result, the animal
breaks through the tough water-surface lm
to become airborne. The passage through the
surface lm robs them of almost 90 percent of the energy
they have just generated; nonetheless, once free of the
water their velocity can reach 2.16 feet (660 mm, about
220 body lengths) per second, and they can travel as far
as 6.7 inches (170 mm, or 57 body lengths), although
the average distance is more on the order of 3 inches (80
mm, or 27 body lengths). This isnt smooth ight! Seat
belts denitely need to be fastened and the tray tables had
better be upright and uptight as the animals spins, ass
over teakettle, somersaulting at the rate of about 7,500
rpm. This is violently awesome behavior for these little
animals, but as a predator avoidance maneuver, it works.
The average distance in the air is more than three times
the length of the pursuing sh, and further than the sh
can typically see. In one study of 89 sh attacks causing
copepods to y up, up, and away, only one of those was
captured and eaten by the same sh that had been pursu-
ing it. Again converting these values to human equiva-
lents, it would be as if a 6-foot-tall human swimmer were
to make one powerful breast stroke and be propelled from
the water to y about 350 feet in about a quarter of sec-
ond. Of course, the landing might be a bit diceytiming
the approximately 1,875 somersaults so that the landing
was appropriately graceful and awe-inspiring would take
more than a bit of practice. A human that could success-
fully emulate a copepod could go faster than a speeding
pistol bullet, and surely leap a tall building with a single
bound. One might wonder, though, if Lois Lane would
nd all those thoracic appendages appealing. Depending
A relatively large calanoid has been measured
moving at about 420 body lengths per second.
upon the size of the animal, copepod muscles range from
single cells (small animals) to extensive muscle masses
(larger species). The mandibles and thoracic appendages
are often driven by muscles masses that are quite large
relative to the size of the animal.
It is one of the more interesting animal paradoxes that,
while crustaceans are often vastly different in appear-
ance and behavior, those differences are largely due to
variations in the external morphology, particularly the
appendages. The guts are surprisingly consistent across
many groups of arthropods. It is as if natural selection
has acted to provide as many exterior packages as pos-
sible to put the same guts into different situations. Of
course, it is entirely possible that all the previous state-
ment really means is that we havent looked at the inter-
nal anatomy of enough small bugs, and that we are not
sophisticated enough to discern the phenomenal differ-
ences between them. Nonetheless, as one might expect
by now, the copepods denitely excel in some aspects of
their internal anatomy.
The gut in most copepods is a simple J-shaped tube,
which starts at the ventral, rear-facing mouth and termi-
nates in the anus on the top of the next-to-last abdomi-
nal segment. The esophagus passes anteriorly toward the
head end of the animal and then passes vertically, often
expanding into a bag-like region in the head. After this,
the gut passes to the rear and forms the large stomach.
Midgut glands found on either side behind this region
secrete digestive enzymes into the stomach. A short,
straight intestine exits the stomach and merges with the
hindgut in the anal segment in the abdomen.
Food is ground to small fragments by the mandibles.
As the food passes out of the mouth cavity into the
esophagus, a thin, cellophane-like, chitinous, peritrophic
membrane is secreted around each blob or bolus of food.
This peritrophic membrane is permeable to digestive en-
zymes and digested food, but it protects the gut wall by
preventing the gut contents, which might include abra-
sive materials like ground silica diatom shells, from ac-
tually contacting the tissues of the gut wall. The mixing
of ground-up food and digestive enzymes occurs in the
esophagus and anterior gut. Digestion takes place in the
stomach, with absorption typically occurring in the pos-
terior part of the stomach and the midgut/intestine. The
hindgut compacts the food boluses into fecal pellets that
are expelled from the anus.
Reproduction in copepods follows a basic pattern that is
surprisingly similar throughout the group. During mat-
ing, the male glues a packet of sperm to the female near
or on her gonopore. The sperm are amoeboid and crawl
into the female. Fertilization may be immediate or de-
layed for several weeks. Eventually the eggs develop and,
enclosed in an eggshell, are extruded from the female. In
calanoids they are shed into the water, but in the other
groups they are held in an external egg sac. In the latter
case, they are usually carried long enough for the em-
bryo to develop to the rst free-living stage, the nauplius,
which is released when the eggs hatch.
The nauplius is the fundamental larva of crusta-
ceans. Almost every crustacean either passes through
a free-swimming naupliar stage, or a naupliar stage is
recognizable in their development to some other hatch-
ing stage. The rst nauplius is a small, unsegmented,
teardrop-shaped animal with three pairs of appendages,
referred to by their adult names of rst and second an-
tennae and mandibles. The mouth, covered with a la-
brum, is located between the mandibles, and the short
gut terminates in the anus at the end of the body.
Copepods pass through a total of six naupliar stages.
As they grow they molt into the next stage. With each
molt, the appendages and the body can be remodeled
or new appendages can be added. The sixth naupliar
stage molts into a second larval type called a copepodid
larva, wherein the thorax and abdomen become dis-
tinctly separated. In the rst copepodid larva, there may
be only three thoracic appendages, and the abdomen is
still unsegmented. There are ve copepodid stages, each
of which becomes progressively more like the adult. The
terminal, or last, molt results in a sexually mature adult.
In most calanoids, the development from egg-laying
to reproductive adult takes about a year, and the ani-
mal typically has a one- to two-year life span. In many
harpacticoids and other smaller species, the whole se-
quence can be accomplished in less than a week, and the
life span is correspondingly shorter.
Copepods are almost absurdly common in the oceans,
but are surprisingly lacking or unknown in marine
aquaria. Free-living harpacticoids often enter the sys-
tem with live rock; providing the system has some suit-
able microhabitats, they will persist indenitely. Other
harpacticoids may be common parasitic pests on corals,
and still others enter the aquarium when live cultures
are added as food. In many cases these species do not
persist, as they are eaten by one of the various appropri-
ate predatorswhich is why they were added to the tank
in the rst place.
For aquarists with some means of magnication,
examination of the copepods found in their tanks may
become an interesting and rewarding experience, and a
good adjunct to the rest of their hobbyist experience. For
other aquarists, the best that can be said for copepods is
that they are the preferred and natural food for many,
particularly SPS, corals. And that fact is probably suf-
cient to ensure that copepods, or copepod mimicssuch
as brine shrimp nauplii, Artemia speciesshould be add-
ed to the tank with regularity and in abundance.
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ith the explosive growth of global aquaculture in this new century,
it is sometimes forgotten that the farming of aquatic organisms is
an ancient art, believed by some to have originated in China with the
breeding and rearing of domesticated carp almost 4,000 years ago.
Over the years, the feeding and rearing of fry and larvae have pre-
sented major challenges to the successful culture of many marine shes
and invertebrates, and using Artemia nauplii (baby brine shrimp) and rotifers as
rst foods has brought modern aquaculture to its present state. Although the one-
two, rotifer-Artemia punch, as used in clownsh culture, is capable of raising many
organisms, it has its limitations. Many experimental breeders have discovered in
recent decades that the calculated use of copepods has the potential to allow aqua-
culture breakthroughs with ever more difcult species.
Laser scanning
with enhanced
colors shows
the mouth parts
of Centropages
hamatus, designed
to crush the shells
of diatoms. Image
by Jan Michels,
University of Kiel,
by Eri k Stenn
Practical pods










Copepods have been found in gut content analyses of numerous larval shes. Nutritional stud-
ies have shown that copepods have the proper composition and the magical correct ratio of
fatty acids (DHA:EPA) for the development of marine larvae. Some speculate that copepods are
responsible for delivering probiotics to the predators that eat them or that they harbor enzymes
critical to the development of larval shes and invertebrates.
Early work looked at most of these topics and also focused on harpacticoids such as Tisbe,
Nitokra, and Tigriopus. Fish larvae fed copepods almost always demonstrated higher survivals
and/or fewer deformities than those fed traditional rotifer/Artemia diets. Copepods, in general,
are known to feed on bacteria, phytoplankton, fecal matter, detritus, other copepods, sh lar-
Above: Nannochloropsis
phytoplankton culturing
columns at AlgaGen.
Below, left: an
unidentifed species
cultured by AlgaGen;
right: Tangerine
ReefPod with eggs.
vae, and other organisms from the planktos. Their food gets converted into essential fatty acids
that are consumed by a host of other marine organisms, many of whom cannot get them any
other way and need them to survive.
To the aquarist, copepods have value as a food source for corals, shes, and other pod life.
Copepods that eat detritus can actually help keep an aquarium system clean because they eat
leftover sh food, feces, and plant debris, and then convert that waste into fatty acidcontain-
ing copepod biomass that is then reintroduced into the food chain. In nature there is ample
space and resources for copepods to thrive, but in the reef tank it is a different matter. Due to
animal density and lack of food, copepod populations in a reef tank can become exhausted fairly
quickly. It is important to repeatedly add copepods to a system because they are constantly being
consumed by amphipods, wrasses, corals, Anthias spp., mandarins, pipeshes, and so on.
There are numerous orders of copepod, and not all are ideal to have in a reef tank or in
culture. In selecting copepods to add to a tank, harpacticoids (order harpacticoidea, benthic co-
pepods) are a good choice. They generally survive well in a reef tank because they have access to
food. They can tolerate pumps, skimmers, lter material, and ow, and will persist and poten-
tially bloom (depending on the species
and how the tank is stocked). Harpacti-
coids are a good food source for numer-
ous tank inhabitants: corals, shes, and
other pods.
The order calanoidea (pelagic or free-
swimming copepods) can persist in a reef
system but probably will not bloom, due
to the lack of available food. Ironically,
divers tell stories of swarms of calanoids
so thick during the dusk/dawn periods
that they cannot see their hands in front
of their faces. It is assumed that the
swarms of calanoids and cyclopoids that
appear on the reefs are the copepods re-
sponsible for feeding the corals and lar-
val shes en masse. The primary food for
many of the pelagic copepods is phyto-
plankton (unicellular planktonic algae).
In nature, phytoplankton is present in
sufcient quantities to at least tint the
water slightly brownsometimes very
brownand provide enough food for
the copepods. However, the operation
of a typical reef tank aspires to a differ -
ent aestheticcrystal-clear waterwhich
means not enough food to sustain a
healthy calanoid population. Still, calanoids are a very important component of the food chain,
and adding them to a reef tank creates health benets for the tanks inhabitants.
In addition to a being an important component of an aquarium ecosystem, copepods have been
identied as a critical rst food in the breeding of numerous edible and ornamental shes. The
important aspects of copepods are:
- Mutritional comosition and valuc: a crcct ratio o atty acids (OHA:EFA) lus a con-
jectured host of other benecial nutrients. May reside for a longer time in the gut.
- Sizc: tlc naulii (carly stagc cocod) can bc small cnougl or a larval sl to ingcst.
- Movcmcnt: tlc movcmcnt o a cocod is sastic, }crky, sto-and-go, wlicl causcs tlc
most incredible (and apparently involuntary) feeding response from shes.
the smallest
available calanoid
copepod, is a
fsh feed utilized
in cutting-
protocols for
reef basslets and



















Arguments against the use of copepods include:
- Sacc rcquircmcnts
- "Low" roduction dcnsitics
- Escac rcsonsc by tlc naulii, }uvcnilcs
- Mccd or livc lytolankton
In a commercial aquaculture facility, these arguments can be the decisive factors in the
choice to rely on wild plankton tows instead of producing copepods in-house. Wild plankton
tows are a gamble because in addition to capturing copepods and other potentially useful food
items, they can also introduce unwanted organisms, viruses, and diseases. In addition, the use
of cultured zooplankton closes the loop on sustainability, a direction the hobby is moving in.
In the marine hobby it may be critical to raise copepods because of the market price of the sh
species being cultivated and the fact that many valuable marine ornamentals can only be reared
with copepods.
Admittedly, the intensive use of copepods is new. Marine breeders often believe that there is a
recipe for matching rst foods with each species of sh, but we are far from having this complicat-
ed question all gured out. It may be one copepod for a specic group of shes; it may be a specic
copepod for each stage of development for another group
or species of sh; it may be different for each species un-
dertaken; it may be that we can take four or ve different
copepod species and make them work for numerous spe-
cies of shes, as we have done with rotifers and Artemia.
1odd Gardncr at tlc Long Island Aquarium lad suc-
cess feeding rare reef basslets (Liopropoma spp.) and
grammas (Lipogramma spp.) Parvocalanus, a small co-
pepod, then transitioning them on to Acartia tonsa as
the larvae developed. Todd found that the Parvocalanus
nauplii had an escape response he called hyperspac-
ingthey would move from one side of the tank to the
other in a ash, too fast for the larvae to catch them.
And the Parvocalanus population grew so fast that they
overwhelmed his production system. He nally found
success with his Candy Basslets using Parvocalanus in
conjunction with Acartia tonsa. When one or the other
was not present, he was unable to rear his larvae.
Frank Baensch has reported that Centropyge dwarf
angelsh species can be reared on Parvocalanus, but the
larvae do not require such small nauplii and do better in
the presence of other copepods. His feeling is that the
later stages of the Parvocalanus nauplii are not an ideal
food source for older larvae; for these he recommends
Oithona and Apocyclops.
Mattlcw L. Wittcnricl, Fl.O., Eric Cassiano, and
tlc rcscarcl grou at tlc 1roical Aquaculturc Lab in
Ruskin, Florida, have isolated a small cyclopoid, Oithona
colcarva, witl wlicl tlcy lavc lad succcss. [ason Lcmus,
Ph.D., has had great success using Acartia tonsa in his
work with snappers at the Southern Mississippi Gulf
Coast Rcscarcl Laboratory. Wc lavc lad succcss at Al-
gaGen using Pseudodiaptomus (a large copepod) in rais-
ing carangidae larvac (Lookdowns, Falomcta) as wcll as
Reidii seahorses.
Wittenrich speculates that there may be a specic
food source (or combination of sources) for each sh
species, and that this may be linked to the ecological
pelagicus, a
calanoid copepod
whose small nauplii
are suitable for
many species of fsh
Bottom: Acartia
tonsa, a calanoid
that produces
small nauplii used
for flter-feeding
corals, and many
larval fshes.
Below: Tisbe
biminiensis, small
harpacticoid with
egg mass.
Bottom: 1-gallon
copepod culture
niche of that sh. He points out that the diet of larval shes can often be generalized at the fam-
ily level, but just like the adults, larvae can be categorized as generalist (dinoagellates, tintinids,
and perhaps large diatoms) or selective feeders. A clownsh larva, for example, will consume
most zooplankton organisms placed in front of it, whereas many other damselsh larvae will
select a single species and stage of cyclopoid copepod.
It is important to note that the copepod is not, by itself, the magic bullet, but goes hand in
hand with technique. There is much we dont know, and many questions face us: Is it important
to keep searching for the correct species of copepod for each individual sh species we are at-
tempting to breed? What are we looking for? Does the
nutritional content of a copepod vary based on its life
stage? A point to consider is that the use of copepods
as an initial feed may only be necessary for a nite
period of timefor instance, for a few days at a criti-
cal juncture in the larval development. If this were
the case, it would not be necessary to set up a large
copepod production system, since there are commer-
cial sources for quantities of the right copepods being
produced for on-demand, just-in-time orders.
At AlgaGen, we have selected and market a handful
of species that have proven useful in reef aquarium
and breeding applications. They have also proven to
be reasonably easy to culture.
Tisbe biminiensis
Tisbe biminiensis is a small harpacticoid copepod that lives on aquarium surfaces and swims in
the water column. It reproduces sexually, with 10 to 70 eggs deposited in an egg sac attached
to the females genital segment. Nauplii are 55140 micrometers wide, and undergo six nauplii
stages and ve copepidid stages before becoming sexually reproducing adults at an age of 912
days. Their natural lifespan is roughly 1520 days.
T. biminiensis eat detritus and microalgae in the aquarium. Their small nauplii are an ex-
cellent food for aquarium lter feeders and sh larvae. The adults are eaten by small bottom-
feeding shes such as gobies, dragonets, blennies, and juvenile seahorses. T. biminiensis is a great
choice to stock into refugiums to perform tank hygiene control and provide background live
feeds to the main aquarium.
The culture preferences for Tisbe biminiensis arc: Liglt to mcdium acration; tcmcraturc 22-
30C; Salinity 20-35ppt; feed phytoplankton, such as PhycoPure CopePod Blend
Ventral anatomy,
greatly enlarged
and artifcially
of Temora
Image by Dr.
Jan Michel,
University of Kiel,
Acartia tonsa
Acartia tonsa is a copepod thats been around the block for a while now, but generally not in
commercially available quantities. A. tonsa is a small to medium (1 to 1.2 mm) calanoid co-
pepod that lives in the water column throughout its life. A. tonsa is known to shed eggs which
will sink to the substrate; these hatch out as 65-120 micrometer wide nauplii which undergo 6
nauplii and 6 copepodid life stages before reaching the adult phase after 9-12 days. Adults will
live for 20 to 25 days.
A. tonsa is an opportunist planktivore that will consume colloidal particles, detritus, micro-
algae and zooplankton. This broad diet may be the reason A. tonsa is one of the easier calanoid
copepods to culture, but it should be cultured in low densities to reduce cannibalization of the
nauplii by the adults. The nauplii of Acartia tonsa have been shown to be viable rst foods for
numerous marine sh larvae including Chrysiptera sp. damselsh; Dr. Andrew Rhyne demon-
strated a low-intensity growout system for dragonet larvae (Synchiropus sp.) using A. tonsa.
Copepod culture is not a difcult endeavor and is only about available time and space. For small-scale home
production, the equipment needed consists of appropriate containers (two or three one-gallon jars with
lids), rigid air rod, air pump, gang valve to control air ow and 2 screens (55um, 150um; screen size will
vary depending on the species cultured), an area that can maintain constant temperatures of roughly 75-85F
dccnding on sccics. Liglt may not bc ncccssary, but wc likc diusc liglt.
We rinse out the containers several times with saltwater prior to use to remove any production residues.
Either test the water of the incoming culture or ask for salinity parameters; match them if possible. If cultur-
ing calanoids we will stock a one-gallon container with 1,000-2,000 reproductive copepods. Add enough live
microalgae (T. Isochrysis galbana) to tinge the water with a light tea color. Feeding less but more frequently is
preferred. Aeration of one bubble per 2-3 seconds is plenty; aim for large bubbles, very gentle aeration. As a
general statement, calanoid copepod nauplii (babies) are reproductive after 8-9 days. It is important to sepa-
rate the nauplii from the adults for several reasons; the presence of nauplii creates some cue that causes the
adult copepods to stop producing nauplii. In addition, over-population of nauplii can instigate cannibaliza-
tion of the nauplii by the adults. Constantly removing the nauplii from the production system will stimulate
the adults to keep reproducing.
Be mindful of over-feeding. If there is too much detritus in the production system, it will cause fouling
of the copepods and will cause them to sink and die. As a mindless task we recommend changing the water
every week in the following manner. Place the 55um screen in a Rubbermaid tote big enough to keep the
bottom of the screen submerged 2-3 inches (to keep the nauplii copepods from drying out). Place the 150um
on top of the 55um screen and pour the culture through the nested screens. The adults will collect on the
150um and the nauplii will collect on the 55um. Clean out your container, rell and rinse the adults back
into their container. Meanwhile take another container that has been prepared (rinsed and lled with water)
and rinse the nauplii from the screen into that container. Feed both containers lightly and keep aeration at
one bubble per 2-3 seconds. If all goes well your copepod population will grow aggressively and the popula-
tion will incrcasc to numbcrs grcatcr tlan 2-5/mL; now our or ccd out tlc amount ncccssary to gct tlc
oulation back down to 0.5-1/mL. 1lc gcncral liccyclc o calanoids can bc u to 1 montl. By kccing tlc
oulations in cacl }ar maintaincd to 0.5-1/mL tlcrc slould bc good naulii roduction ovcr tlc coursc o
the copepod life span. If there are water changes and size separations performed each week there will always
be an adult container as well as a nauplii container. This approach can be extrapolated to scale.
When raising harpacticoid copepods a similar process can be employed except that one can crush ake
food, use live or processed phytoplankton, or other microparticulate feeds. When using a microparticulate
feed there is a greater chance that water quality parameters will degrade and so water changes are important.
Despite my earlier recommendations, for a few species of harpacticoid or cyclopoid copepods, developing de-
tritus at the bottom of the container is important and a system kept too clean will cause poor performance;
again, this is species specic.
Culture preferences for Acartia tonsa are: Aeration-
light; Temperatures of 24-27C; Salinity of 20-30 ppt;
stocking densities of < 1/ml; feed with live phytoplank-
ton such as T. Iso or PhycoPure CopePod Blend .
Parvocalanus crassirostris
Parvocalanus is the smallest calanoid copepod routine-
ly available. It is a truly pelagic copepod, with nauplii,
copepodids, and adults all living in the water column.
Parvocalanus drops individual eggs, which hatch into 40-
100 micron nauplii that undergo 6 nauplii and 5 cope-
podid life stages before reaching reproductive adulthood
at 6-8 days of age. At only 0.2 to 0.3 mm in size as an
adult, it lives only 12-15 days.
Parvocalanus is recognized by top aquaculturists and
aquarium breeders as one of the best-known live foods to
date for the smallest and most difcult of the marine sh
larvae due to the small size of the nauplii. Parvocalanus is
rich in essential fatty acids and provides good nutrition
for developing larval sh. In aquariums the small size
is optimum for capture by the nest lter feeders such
as feather duster and Christmas tree worms, SPS corals,
even sponges. Parvocalanus populations can expand rap-
idly when optimum culture conditions are maintained.
Culture preference: temperature 24-32C; salinity 25-
35ppt; light aeration; feed live phytoplankton or Phyco-
Pure CopePod Blend .
Pseudodiaptomus pelagicus
Pseudodiamptous pelagicus is a medium-sized (1.2 to 1.5
mm) calanoid copepod that normally lives in the water
column, although adults are noticed for their ability to
attach to surfaces. P. pelagicus males attach to females
for extended periods of time; females carry egg sacs and
release live nauplii which are 75 to 160 microns wide.
These nauplii undergo 5 total nauplii stages and 5 co-
pepodid stages before reaching the adult, reproductive
stage at 9-12 days. They will live for up to 30 days.
P. pelgagicus is a relative newcomer to the aquarium
circuit, and it is proving useful in larval rearing when
rotifers fail. We are just starting to uncover the many po-
tential uses for this species in marine sh breeding. The
adult predisposition for attaching to aquarium surfaces
allows for this species to be stocked into refugiums, al-
lowing adults to continue to produce nauplii to feed the
main tank for extended periods.
Culture preferences: temperatures between 24-27C;
salinity ranging between 20-30 ppt; light to medium aer-
ation; stocking densities of 0.25-1.0/ml; Feed live phyto-
plankton or PhycoPure CopePod Blend .
Unidentified Tangerine Pods
The AlgaGen ReefPod Tangerine is a large calanoid
copepod that we havent been able identify; most likely
they hail from the family Temoridae. This is a big, meaty
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calanoid copepod that spends its life in the water col-
umn. The female carries egg sacs that hatch out nauplii
that are 100-200 microns wide. The nauplii reach repro-
ductive adulthood at 12 to 20 days and 2 to 2.5 mm in
length; theyll live for up to 40 days.
Tangerine copepods will eat small suspended detritus
particles and microalgae in the aquarium. They produce
large nauplii, which are an excellent food for aquarium
lter feeders and sh larvae. The adults are eaten by nu-
merous water-column feeding sh such as clowns, dwarf
angels, anthias, and seahorses, to name just a few.
Culture preferences: Temperatures between 22-25C;
Salinity o 15-30t; Liglt to mcdium acration; Fccd livc
phytoplankton or PhycoPure CopePod Blend .
There is one other noteworthy copepod readily available
to aquarists at this time; Tigriopus californicus. This spe-
cies is commonly recognized by the Reed Mariculture
trademark name Tigger-Pods.
Tigriopus californicus is a harpacticoid copepod with a
predisposition for active swimming in the water column.
It is generally 1 to 1.5 mm in size as an adult. Females
can carry anywhere from 13 to 31 eggs per clutch. Nau-
plii go through 6 nauplii stages and 6 copepodid stages
before reaching reproductive adulthood at 20-30 days.
Tigriopus californicus have proven themselves to be
any easy copepod to culture, perhaps owing to the ex-
ceptional diversity of their natural habitats all along
the western coast of the United States.
Reed Maricultures culture instruc-
tions include the use of simple aeration
and feeding with prepared algae-based
feeds (such as Reed Maricultures Phyto
Feast), otherwise cultured at room
temperatures with regular strength salt-
At AlgaGen we have been producing
copepods for over 10 years at various
scales. The most basic production occurs
in 5-gallon containers but we also pro-
duce in 1,000-gallon systems. The tech-
niques described above are basically how
we approach production. However, some
aspects are automated. Mick Payne has
done some great pioneering work with
production and production systems and
his prior work could serve as a platform
to develop your own system. There is a
nice review article in Aquaculture 315
(2011) p. 155166 that discusses sys-
tems and species.
We are very fortunate to work near-
by Florida's Indian Rivcr Lagoon and
warmer Atlantic Ocean waters that give
us a huge source to tap for new species.
When possible we perform plankton
tows or get tows from friends who are
out on the water. The samples are diluted
and examined in a petri dish under the
microscope. Painstakingly, individuals
arc isolatcd and laccd in 2-L contain-
ers, fed phytoplankton, and watched.
If and when something is reproducing
well, seems size-suitable and practical to
culture, we send a sample off to be iden-
tied. For all of our isolates, species are
maintaincd in 2-L }ars tlat arc cd and
thinned out weekly. Once a culture has

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been identied as being useful, we send samples out to
collaborators to get their feedback on handling, larval
survivals, etc.
Copepods are a keystone group in marine food webs in
the wild, and their importance in aquaculture is likely
to grow exponentially. Over the past years cultured co-
pepods have become much more easy to nd and read-
ily available to anyone interested in obtaining them. The
importance of copepod availability has been identied
as a crucial service for aquaculturists, and even home
breeders can nd numerous sources for copepod cul-
tures. As more pod species are found and as people
share their experiences feeding challenging larval shes,
we can hope to hear of more exciting aquaculture break-
throughs and much expanded availability of marine
aquarium organisms.
Erik Stenn is AlgaGens founder and chief scientist, based in
Vero Beach, Florida.
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obtained my rst aquarium at the age of nine years
and became infected with the aquatic virus, a pas-
sion that has never left me. After accumulating ex-
perience with the keeping and breeding of discus, in
1985 I graduated to the marine aquarium hobby with
a 132-gallon (500-L) tank. While diving in the Mal-
dives in the same year, my desire to add colorful
stony corals to my captive reef was born.
Back then, however, keeping stony corals was
considered almost impossible. The reef aquarium
hobby was still in its infancy, and I experienced the
(expensive) development of stony coral aquarium
technology rst hand. One by one, I upgraded the
lighting, the current pumps, the skimmer, and the
calcium reactor with new and better products.
The actual realization of my stony coral dreams
came at the beginning of the new millennium, when
Thomas Pohl developed the Zeovit system. This per-
mitted the keeping of corals in a nutrient-poor environ-
ment, using ltration over zeolites and supplementation
with small doses of bacteria, nutrients, and trace ele-
ments. At last I had what I had always wanted: lots of
These small-polyp stony corals display the pastel colors typical
of the Zeovit system. Note Tridacna gigas mantle, foreground.
A gigantic clam
in my living room
aquarium portrait | BY WI LLI WEHNER
Front view of the aquarium
at eight years of age.
Achieving a shell length of 2 feet (60 cm) in just seven years, my once-tiny Tridacna gigas
currently occupies about a third of its 6.5-foot (2-m) aquarium. Part of the reef structure
has already been removed to make room for itand its still growing!
well-fed shes swimming among colorful stony and soft
corals. I always used my favorite diving area, the Red Sea,
as my model.
After 20 years, algae had eaten deep into the white sili-
cone of my aquarium. I had to deal with this in order
to prevent the looming catastrophe of a leaking tank.
In the course of a four-week campaign I renovated the
living room and had the new 6.5-foot (2-m) tank con-
structed on the spot. After a maturation phase of only
two weeks with 12 live pieces of tufa rock from my rst
reef aquarium, I transferred the shes and corals without
any complications or losses.
I had always been attracted to colorful Tridacna. In spring
2005 I added a 3-inch (8-cm) T. gigas that I bought from
star of
the reef
aquarium is
this massive
Giant Clam
gigas), whose
shell is
already 2 feet
(60 cm) long.

SIZE, VOLUME, TIME IN OPERATION: 79 x 32 x 24 inches
(200 x 80 x 60 cm); 254 gallons (960 L); Eight
Acropora, Montipora, Pavona, Seriatopora, Stylophora,
Turbinaria. LPS: various Acanthastrea, Blastomussa,
Caulastrea, Euphyllia, Fungia, Mesophyllia, Plerogyra,
Trachyphyllia, Duncanopsammia axifuga.
pitularia, Litophyton, Sarcophyton, Sinularia.
OTHER INVERTEBRATES: 2 Tridacna crocea, T. derasa,
T. gigas; brittle stars, reef lobsters, various hermit
crabs and gastropods, 3 Entacmaea quadricolor.
FISHES: 2 Amphiprion bicinctus, 2 Calloplesiops
altivelis, 11 Chromis viridis, 2 Gobiodon rivulatus,
Pomacanthus navarchus, 13 Pseudanthias squamipin-
nis, Pseudocheilinus hexataenia, Salarias ramosus, 3
Sphaeramia nematoptera, 2 Zebrasoma avescens, Z.
DECOR: Exclusively tufa rock (already used for 20
years in the previous aquarium and well colonized
with sponges and small organisms), aragonite sand
with a little crushed coral and zeolite as substrate.
LIGHTING: Three 250 W HQI (lamps: 10,000 Kelvin
BLV-HIT) + two 80-watt T5 lamps (Korallen-Zucht
Super Blue); lighting period 10 hours HQI, 12
hours T5; 9-watt moonlight.
WATER MOVEMENT: 4 Tunze Streams, partly controlled
by Single-Controller and moved via Osci-Motion;
reduction at night; maximum total pump output
10,567 gallons (40,000 L)/h.
WATER MANAGEMENT: Bubble King 250 internal pro-
tein skimmer, permanent ltration over .8 gallons
(2 L) zeolite + .2 gallons (.8 L) activated carbon.
reactor; addition of all products from Korallen-
Zucht; 16-gal. (60-L) partial water change weekly.
PRECAUTIONS: Tunze Nanostream (output: 1.321
gallons [5,000 L]/h) connected to a backup power
supply with three car batteries; cooler from Aqua
Medic for the summer months.
OWNER: Willi Wehner, Trogen bei Hof, Bavaria.
Thomas Pohl in Coburg to the Tridacna crocea and T. de-
rasa already in my aquarium. Since then, the T. gigas has
grown a lot. The prerequisites for this are optimal water
quality and maintenance, thanks to which my SPS cor-
als are also colorful and thriving. Because of their vigor-
ous growth I am regularly able to pass
on cuttings of soft and stony corals to
other aquarists and aquarium stores.
In the eight years that the new tank has
been set up, there have been only oc-
casional minor problems. Slime algae
(cyanobacteria) have appeared twice
and then disappeared after a while. The
trick is to wait quietly and patiently
without rushing into measures that,
in the worst-case scenario, could make
things even worse. But when faced
with a mass proliferation of little red
atworms (turbellarians), I added a
Six-Stripe Wrasse and a pair of man-
darinshes, who rapidly got the plague under control.
At times, Aiptasia have proliferated in the tank due
to the generous feeding of my more than 40 shes. For
years they were decimated by a Pacic Blue Tang (Para-
canthurus hepatus), which had an undoubtedly species-
Other actinians, like this splendidly
colored Bubble-Tip Anemone (Entacmaea
quadricolor), home to a pair of Amphiprion
bicinctus, are also doing well.
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The world is yours.
The earth is composed of water71.1% to be exact. But when it comes to
tropical sh, weve really got it covered. Not only with exotic varieties from
around the globe, but with the highest level of quality, selection and vitality.
Ask your local sh supplier for the best, ask for Segrest.
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atypical feeding preference
for these unwelcome guests.
After the tang died after 10
years in my aquarium, the
Aiptasia proliferated wild-
ly, so in November 2011 I
added a Majestic Angelsh
(Pomacanthus navarchus),
which eliminated the prob-
lem within a few weeks. So
far it hasnt nibbled at corals
or clams.
I very much hope that my
Tridacna gigas doesnt reach
the maximum size of 4.25
feet (130 cm) cited in the literatureif it does, I will have
problems. But hopefully it will live to be at least as old
as the oldest sh I currently have, an 18-year-old Yellow
Tang (Zebrasoma avescens).
Videos of my reef aquarium and my discus tank can be seen via
the following link:
Thanks to top-quality equipment
and painstaking maintenance,
there is no confict between
nutrient-poor water for the stony
corals and well-fed fshes.
Ask Your Local Specialty Retailer, or
Buy Direct Online: www.fritzpet.com/reef-chemicals
Reef Chemistry Trusted by Professionals, Now In Hobbyist Sizes
Two Little Fishies
We didnt think so. Thats something you probably like doing yourself, and its really a
pleasure when you have the right tools to do the job. Thats where we come in. Two Little
Fishies AquaStik Coralline Red and Stone Grey are underwater epoxy putties with
clay-like consistency for easy attachment of corals. The colors of AquaStik blend perfectly
with rock for a natural effect. Both colors are available in 2oz and 4oz sizes.
CorAfx is an ethyl cyanoacrylate bonding compound with viscosity similar to honey.
Use it for attaching stony corals, gorgonians, and other sessile invertebrates in natural
positions on live rock, or use in combination with AquaStik to attach larger coral heads.
CorAfx Gel and CorAfx Pro cyanoacrylate bonding compounds have a thick gel
consistency that makes them very easy to use for attaching frags of stony corals, zoanthids,
and some soft corals to plugs or bases. AquaStik, CorAfx, CorAfx Gel and CorAfx Pro
work on dry, damp, or wet surfaces, cure underwater, and are non-toxic to sh, plants and
invertebrates. Developed by aquarium expert Julian Sprung.
Would you like us to showyou where to stick it?
Genicanthus semifasciatus rarely turns up in the aquari-
um trade, but the tiger-striped males are spectacular. If
possible, it should be kept in pairs or groups with just
one male in the aquarium.
This species is basically quite hardy and usually feeds
readily, something that should be checked by a test feed-
ing prior to purchase. The minimum aquarium size is 75
gallons (285 L), but signicantly larger is best. Like other
Genicanthus species, in most cases it can be housed with
sessile invertebrates without problems.
The Japanese Swallow Angelsh occurs in the western
Pacic from southern Japan to the northern Philippines.
Genicanthus semifasciatus grows to around 8.25 inches
(21 cm) long and is broadly similar in body form to oth-
er Genicanthus species known in the aquarium hobby.
This species exhibits pronounced sexual dimorphism;
the males and females are very differently colored.
species spotlight | DANIEL KNOP
Genicanthus semifasciatus
Phylum: Chordata (vertebrates)
Class: Osteichthyes (bony shes)
Order: Perciformes (perch-like shes)
Family: Pomacanthidae (angelshes)
Genus/species: Genicanthus semifasciatus
Japanese Swallow Angelsh








Head shots of Japanese
Swallow Angelfsh (Genicanthus
semifasciatus) female (left) and
male (right).
Japanese Swallowtail Angelfsh
(Genicanthus semifasciatus), male.
Japanese Swallow Angelfsh
(Genicanthus semifasciatus), female
(male in background).
Males have distinctive vertical dark bands on a sil-
very background on the upper half of the body, and the
lower half of the body is light silvery. A bright golden-
yellow wedge extends from the head (mouth to fore-
head) to the center of the body, and the black bands and
yellow areas intensify during courtship. Females have a
yellowish dorsum and a silvery white underside, a typical
black forehead mask with two vertical bars, and a black
caudal-peduncle band.
In 1970 the female of this species was erroneously sci-
entically described as Holacanthus fucosus. It was regard-
ed as a separate species until 1975, when the pronounced
change of color accompanying the female to male sex
change was observed. In the wild, it lives in large groups
in which females outnumber males or in small harems
consisting of one male and several females. Females will
transform into males if the opportunity presents itself.
The Japanese Swallow Angelsh lives in coral-covered or
rocky outer reef slopes at depths of between 50 and 660
feet (15200 m), sometimes even deeper.
Allen, Gerald R., R. Steene, and M. Allen. 1998. A Guide to
Angelfshes & Butterfyfshes. Odyssey Publishing, Perth,
Debelius, Helmut and R. Kuiter. 2006. World Atlas of Marine
Fishes. Hollywood Import & Export, Gainesville, Florida.
Lieske, Ewald and R. Myers. 1998. Coral Reef Fishes. Princeton
University Press, Princeton, New Jersey.
Michael, Scott W. 2004. Angelfshes & Butterfyfshes, Reef Fishes
Book 3. Microcosm/TFH, Neptune City, New Jersey.
The mind has exactly the same power as the hands: not merely to
grasp the world, but to change it. - Colin Wilson
Our worst fear is not that we are inadequate, our deepest fear is that
we are powerful beyond measure. - Nelson Mandela
The miracle, or the power, that elevates the few is to be found in
their industry, application, and perseverance under the prompting of a
brave, determined spirit. - Mark Twain
Knowledge is power. - Francis Bacon
What is Power?
This is Goniopower

Justin Credabel

f or novi ces i n the mari ne aquari um hobbyDANI EL KNOP
irtually no other type of aquarium reacts as
badly to heavy metals as a reef system. But in
addition to visible metal components on or in
the aquarium that may come into contact with
salt water, there are hidden metals that can cause
the concentration of certain metal ions in the
water to rise to toxic levels.
This feature usually deals with the setting
up and maintenance of a healthy marine aquar-
ium and how to avoid problems. In this issue we
would like to take a different route and discuss a
problem that may already be present in your aquari-
um: heavy-metal poisoning. Some marine aquarists,
even veterans, are unaware of how dangerous even
a slightly elevated heavy-metal concentration can be
in this type of aquarium. Invertebrates are extremely
sensitive to these metallic substances. They are prac-
tically always present in natural seawater in very low
concentrations, at which many of them are also im-
portant for certain physiological processes. But if
the concentration increases, some metabolic pro-
cesses may be disrupted, or there may be full-blown
poisoning that can seriously harm invertebrates.
Suspect heavy-metal poisoning in aquariums that are
doing poorly for no apparent reason, or aquariums in
which corals arent thriving and nuisance algae are pro-
liferating in the absence of elevated phosphate and ni-
trate concentrations that could explain this. Even more
Heavy-metal poisoning
in the aquarium, part 1
Only by preventing
harmful metal
ions from tainting
the water can
one maintain a
fascinating reef
aquarium like this
one belonging to
Matthias Paul. If a
problem occurs,
all the aquarium
equipment should
be examined in
detail. Above is
the flter tank in
Krzysztof Trycs

suspicious are the occasional deaths of mollusks,
such as gastropods, or echinoderms, such as star-
shes or sea urchins. And a puzzling shrimp die-off
can also point to heavy metal contamination. Admit-
tedly, other factors could be responsible, but heavy-
metal poisoning should be ruled out as far as possible
through rigorous checking.
First of all, carefully consider the aquarium, all
its equipment, and any equipment used in mainte-
nance, even if it only briey comes into contact with
the tank water. Stainless steel forceps can release metal
ions in seawater. Glass-cleaning razor blades should be
used only as long as they show no signs of corrosion, in-
cluding in the non-visible area beneath a plastic holder.
Metal hose ttings are fundamentally problematic in
the marine aquarium hobby. They can even cause harm
without direct contact with water, for example through
salt encrustation and periodic condensation that ulti-
mately causes metal ions to enter the aquarium water
unnoticed. And defective plastic encapsulation of metal
components, for example pump impellers, can poison
the water. Any place where metals are usually unavoid-
able in the marine aquarium should be monitored care-
fully in order to discover any water contact or corrosion.
If your search doesnt turn up anything obvious,
check places where you wouldnt normally expect to nd
any metal. There are three common culprits: rst, the
rockwork. Live rock and faux rock are harmless as a rule,
but some limestones originate from earlier geological ep-
ochs, for example from a limestone quarry. Such rock
may contain metal ores hidden in the interior. If this is
the case, you will become aware of it only when you have
serious problems in the aquarium for which no alterna-
tive cause can be discovered.
The second suspect is the substrate. Anyone who uses
materials such as foraminiferan sand, live sand, or lime-
stone granulate runs the risk of introducing hidden met-
als into the aquarium. If you use coral sand and notice
that individual grains of substrate stick to the magnetic
glass-cleaner, there could be iron ore in the sand. It is
unclear whether this can be attributed to impurities that
get in during processing (cleaning, grading, packing)
in the country of origin or whether individual grains of
the coral sand actually contain iron. Thorough check-
ing with a magnetic glass-cleaner or other strong magnet
can exclude this problem. The nal possibility is a net
lter bag supplied with a phosphate adsorber for marine
aquariums. A small Internet mail-order business offered
this product, and its use in some small aquariums led
to inexplicable deterioration of the entire aquar-
ium environment, with stagnating invertebrates,
massively disturbed macro-algae growth, and an
ominous increase in the growth of micro-algae.
Painstaking examination of all details of the
setup eventually revealed a tiny metal spring in
the practical little bag closer that had clearly
undergone a change, virtually undetectable from
outside in normal usage. The closeup shot at left
shows, however, that this was the cause of serious
heavy-metal contamination.
Next time, this section will explain what can be done about exist-
ing heavy-metal poisoning.
Knop, D. 2012. Not so cheap in the long run. CORAL 9 (1): 11314.
Metalliferous particles from the aquarium substrate sticking
to a magnetic glass cleaner. A strong magnet such as
this can be used to screen sand and coral gravel for metal
contaminants or grains of substrate containing iron.
Even a harmless-looking net flter bag
can introduce a tiny amount of metal into
the aquarium system, acting as a sort of
aquarium time bomb. The spring-loaded
bag closure device below shows the
corrosion that developed after submersion
in salt water.
Sales: 970.776.8629 / 877.323.2782 | Email: customerservice@aquariumwatertesting.com
The Worlds FIRST Full Water Testing
Service for Serious Marine Hobbyists!
Silica (Sio2-3)
Natural Seawater Value: 0.040 mg/L
Acceptable Range: 0 0.5 mg/L
Tested Result: 1.5 mg/L
(HIGH) Your silica level is too high. We recommend that you use a silica specifc
R/O membrane in addition to deionization resin for your make-up/top-of water.
You may also use a commercially available phosphate absorber, as these will also
remove some silicate. Silicate is required by many types of sponges for
growth/reproduction, but will also encourage brown diatom algae growth. Any
level above 0.3 mg/L may cause a diatom bloom in the aquarium.
Ammonia (NH3-4) Good 0.010
Nitrite (NO2) Good 0.003
Nitrate (NO3) Good 2.600
Phosphate (PO4) Good 0.020
Silica (Sio2-3) High 1.500
Potassium (K) Low 252
Ionic Calcium (Ca) Good 268
Boron (B) Good 5.200
Molybdenum (Mo) High 0.200
Strontium (Sr) Good 10.70
Magnesium (Mg) Good 1250
Iodine (I) High 0.090
Copper (Cu) High 0.030
Alkalinity (meq/L) Low 2.200
Total Calcium (Ca) Good 397
e recommend that you use a silica specifc
i i f k / f
Phosphate (PO4)
Natural Seawater Value: 0.030mg/L
Acceptable Range: 0 0.25 mg/L
Tested Result: 0.02 mg/L
(GOOD) Your phosphate level is within the recommended range.
We recommend continuing the current maintenance and water
change schedule. The use of a phosphate absorbing resin is
recommended to keep phosphate levels below 0.05 mg/L.

DI STRI BUTI ON: Pacic: Ryuku Islands to Micro-
nesia, south to Australia.
DESCRI PTI ON: With a total length of around
4 inches (10 cm), Neocirrhites armatus is a rather
small hawksh. It is a popular aquarium sh on
account of its bright red coloration and because it is
easy to maintain.
ECOLOGY: As a rule, Flame Hawksh live in pairs
in the wild. They are extremely quiet and sedentary
sh that spend most of their time perching among
sheltering coral branches, usually those of stony
corals of the genera Pocillopora or Stylophora.
nance of a pair of Neocirrhites armatus is possible
even in smaller reef aquariums with a volume of
at least 66 gallons (250 L). These social animals
should not be kept singly, and large corals should
always be present to permit the typical behavior.
They dont necessarily have to be branching stony
corals; for example, mushroom-shaped (Sarcophy-
ton) or nger-like (Sinularia) leather corals are also
accepted as living corals.
FEEDI NG: Flame Hawkshes are not very fussy
eaters, and feed on small crustaceans; once settled
in they will take assorted frozen foods and, usually,
dry food as well.
begi nners l i vestockI NKEN KRAUSE
Flame Hawkfsh
(Neocirrhites armatus)
never fish for a pen again

Sterling Silver
Handmade in Vermont
Fits every Wallets Fold
Look for CORAL Magazine in these outstanding local aquarium shops.
Aquarium Fantasies
340 Eastdale Cir
Montgomery, AL
The Aquarium Shop
2013 Cox Ave
Huntsville, AL
Northside Aquatics
7610 Counts Massie Rd Ste A
Maumelle, AR
Worlds Under Water
2105 Creekview Ste B
Fayetteville, AR
Aqua Touch
12040 North 32nd St
Phoenix, AZ
All Seas Marine, Inc
(distribution only)
1205 Knox St
Torrance, CA
Amazing Aquariums
& Reefs
1842 N Tustin St
Orange, CA
Aqua Exotic
240 Harbor Blvd Ste E
Belmont, CA
Aquarium Concepts
6920 Amador Plaza Rd
Dublin, CA
Aquatic Central
1963 Ocean Ave
San Francisco, CA
Coral Island
1711 W Chapman Ave
Orange, CA
Natural Life Aquarium
131 Southwood Ctr
South San Francisco, CA
Seven Seas
647 Shaw Ave
Clovis, CA
Sierra Saltwater Systems
125 Lassen Rd
Tahoe City, CA
Tongs Tropical Fish
8976 Warner Ave
Fountain Valley, CA
1947 Main St
Watsonville, CA
Whites Pets
5212 North Blackstone
Fresno, CA
Animal Attraction
2518 11th Ave
Greeley, CO
16522 Keystone Blvd, Unit K
Parker, CO
Neptunes Tropical Fish
1970 E County Line Rd Unit A
Highlands Ranch, CO
Aquatic Wildlife Co
179D Deming St
Manchester, CT
House of Fins
99 Bruce Park Ave
Greenwich, CT
Barrier Reef
1921 NW Boca Raton Blvd
Boca Raton, FL
Bio Reef LLC
3653 Regent Blvd #101
Jacksonville, FL
Boardroom Aquatics
12795 Kenwood Lane
Fort Myers, FL
Coral Corral
13510 Prestige Pl
Tampa, FL
Creatures Featured
314 SW Pinckney St
Madison, FL
Eco Reef Corals
2137 S. Tamiami Trail
Venice, FL
Father Fish Aquarium
536 E Venice Ave
Venice, FL
Fishy Business
140 S Ronald Reagan Blvd
Longwood, FL
Ocean Lifers
36037 US Hwy 19 N
Palm Harbor, FL
Orange Park Aquatics
793 Blanding Blvd Ste A
Orange Park, FL
Sea Life Aquarium
& Service
174 Semoran Commerce Pl
Apopka, FL
Ultra Corals Inc
1063 Ingleside Ave
Jacksonville, FL
Aquarium Outtters
175 Old Epps Bridge Rd
Athens, GA
Creation Pet LLC
8265 Hwy 92
Woodstock, GA
Premier Aquatics
1801 Roswell Rd
Marietta, GA
Pure Reef
12900 Hwy 9 North Ste B
Alpharetta, GA
Coral Fish Hawaii
98810 Moanalua Rd
Aiea, HI
Fish, Aquariums & Stu
6112 West Fairview Ave
Boise, ID
Beyond the Reef
205 W Golf Rd
Schaumburg, IL
Chicago Reptile House
14410 John Humphrey Dr
Orland Park, IL
Fish Planet
839 Waukegan Rd
Deerfeld, IL
Sea Escapes
1950 Silver Glen Rd
South Elgin, IL
Sailn Pet Shop
720 S Neil St
Champaign, IL
Inland Aquatics
10 Ohio St
Terre Haute, IN
Aquatic Environments
730 E Kimberly Rd
Davenport, IA
Easy Aquariums
17 A Gorham Industrial Pkwy
Gorham, ME 04038
House of Tropicals
7389F Baltimore
Annapolis Blvd
Glen Burnie, MD
Krystal Clear Aquatics
700 Southbridge St
Auburn, MA
South Coast Scientic
109 McArthur Rd
Swansea, MA
Blue Fish Aquarium
2939 Wilson Ave SW Ste 109
Grandville, MI
Moby Dick Pet Store
3700 Sashabaw Rd
Waterford, MI
7429 S Westnedge Ave
Portage, MI
Oceans and Seas
26085 Gratiot Ave
Roseville, MI
Preuss Pets
1127 N Cedar St
Lansing, MI
16063 Manchester Rd
Ellisville, MO
Gateway Aquatics
4570 Telegraph Rd
Saint Louis, MO
Seascape Studio
3802 S Lindbergh
Saint Louis, MO
Heights Pet Center
895 Main St
Billings, MT
New Hampshire
Aqua Addicts
52 Lowell Rd
Salem, NH
Jays Aquatics
10 Lawrence Rd
Salem, NH
Laconia Pet Center
1343 Union Ave
Laconia, NH
New Jersey
Adams Pet Safari
19 W Main St
Chester, NJ
Aquarium Center
1295 Blackwood Clementon Rd
Clementon, NJ
Ocean Aquarium
6820 Black Horse Pike Rte 40
Egg Harbor Township, NJ
Pets, Pets, Pets
2 JFK Blvd
Somerset, NJ
Tropiquarium & Petland
Ocean Plaza, 1100 State Rte 35
Ocean, NJ
New York
A Reef Creation
4700 Genesee St Ste 112
Cheektowaga, NY
ABC Reefs
527 Charles Ave
Syracuse, NY
Acks Exotic Pets
8107 Brewerton Rd
Cicero, NY
Eddies Aquarium Ctr
1254 New Loudon Rd Rt 9
Cohoes, NY
The Fish Place
141 Robinson St
North Tonawanda, NY
Long Island Aquarium
431 East Main St
Riverhead, NY
Manhattan Aquariums
522 West 37th St
New York, NY
Pet Friendly
845 Manitou Rd
Hilton, NY
Tropical Fish Outlet
2065 Lake Rd
Elmira Heights, NY
North Carolina
Advanced Aquatics
509 Woodlawn Ave
Belmont, NC
Aquarium Outtters
823 South Main St
Wake Forest, NC
Aquatic Consultants
1610 US Highway 70E
New Bern, NC
Blue Ridge Reef & Pet
103 WNC Shopping Ctr Dr
Black Mountain, NC
Croft Pet & Hobby
3800 Reynolds Rd, Suite 200
Winston-Salem, NC
Discount Pet
100 N Main St
Mount Holly, NC
6465 Goldfsh Rd
Kannapolis, NC
Mountains to Sea
14 Sweeten Rd
Asheville, NC
Aquarium Adventure
3632 W Dublin-Granville Rd
Columbus, OH
Belpre Aquarium
1806 Washington Blvd
Belpre, OH
Salty Critter, LLC
4809 Liberty Ave
Vermilion, OH
Saltwater Fanta-Seas
4814 NE 107th Ave
Portland, OR
Daves Aquastock
2301 Duss Ave
Ambridge, PA
Oddball Pets &
262 Joseph St
Pittsburgh, PA
Something Fishy
511 E 21st St
Northampton, PA
The Hidden Reef, Inc
4501 New Falls Rd
Levittown, PA
South Carolina
Aquarium Oddities
1143 E Woodruf Rd
Greenville, SC
Oceans Floor, LLC
179 Halton Rd
Greenville, SC
Sea Critters Depot
3002 Airport Blvd, Ste A
North Myrtle Beach, SC
Austin Aqua-Dome
1604 Fortview Rd
Austin, TX
Birddog & Catsh
115-D Old Boerne Rd
Bulverde, TX
Fish Gallery Houston
2909 Fountain View Dr
Houston, TX
Incredible Pets
1580 Keller Pkwy Ste 50-C
Keller, TX
Pet Advantage
350 Dorset St
S Burlington, VT
Atlantis Aquariums
9602 Patterson Ave
Richmond, VA
11634A Busy St
Richmond, VA
Pet & Aquatic
2408 Wards Rd
Lynchburg, VA
Barrier Reef Aquariums
1717 NE 44th St
Renton, WA
Saltwater City
14150 NE 20th St, Ste F3
Bellevue, WA
West Virginia
Scales & Tails Reptile
& Fish Store
/2 W Washington St
Westover, WV
Reef Wholesale
(distribution only)
12 Vulcan St
Etobicoke, ON
Big Als Aquarium
3511 99th St NW
Edmonton, AB
British Columbia
Paws N Jaws
4750 Rutherford Rd #147
Nanaimo, BC
Progressive Reef
1101790 Island Hwy
Victoria, BC
Red Coral Aquarium
1183604 52nd Ave NW
Calgary, BC
New Brunswick
Maritime Reef
1595 Hickey Rd
St John, NB
Advanced Reef Aquatics
418 Thompson Rd N
Milton, ON
Aquariums by Design
668 Erb St West
Waterloo, ON
Coral Reef Shop
1371 Plains Road East
Burlington, ON
Fish Tail Aquariums
2208 Saint Joseph Blvd #101
Orleans, ON
Living Aquariums
652 Bishop N
Cambridge, ON
Mail Order Pet Supplies
2558 Upper Gage Ave Ste 211
Hamilton, ON
947 Carling Ave
Ottawa, ON
Oakville Reef Gallery
579 Kerr St Unit 2A
Oakville, ON
Sea Life Central
561 Southdale Rd East
London, ON
Raging Reef
10227 Ave Papineau
Montreal, QC
Bayside Corals
501 45 St W
Saskatoon, SK
Pats Pets
1303 Scarth St
Regina, SK
Aqua Blue Distribution
17 Cairns St Unit 4
Loganholme, Queensland
3 Chemin de Maupas
69380 Les Cheres
Water World
Ananda Dutta Lane
Howrah-7111 01
West Bengal
Blue Reefs
82 Triq Guzeppi Mattew Callus
Mosta, Mst 4105
Stunning Corals
Wolvenlaan 285
1216EV Hilversum
South Africa
Aquarium Depot
#1 Mackenzie Park Capital Hill
392 Le Roux Ave
Halfway House 1685
Bioted Marine Ab
Korsgatan 16
434 43 Kungsbacka
United Kingdom
Midland Reefs
Mount Rd Trading Estate
Burntwood, Stafordshire
Email: sales@rvmags.com
CALL (800) 381-1288 | Fax (630) 353-2692
To sell coral
in your store, contact us today:
amoeboid: similar in form to a single-celled
amoeba with an irregular shape and organ-
elles used for propulsion.
arthropod: an invertebrate animal with
an exoskeleton, segmented body, and
jointed appendages. The phylum Arthropoda
includes the insects, crustaceans (shrimp,
barnacles, lobsters, crabs, copepods), spiders
and sea spiders, and others.
cnidarian: member of the animal phylum
Cnidaria, which includes the corals, sea
anemones, sea pens, jellyfsh, and sea wasps.
A characteristic of the phylum is the pres-
ence of stinging-cell cnidocytes, used to
capture prey and in territorial disputes.
cuticle: outermost protective layer in the
exoskeleton of an arthropod.
exoskeleton: in animals, an external skel-
eton that protects the body and provides
support for various body parts. The shell in
crustaceans and mollusks.
integument: in biology, the outer covering
of a bodyits shell, husk, or skin.
motile: in marine biology, an organism that
can move actively and spontaneously, such
as a fsh. The opposite of sessile.
nauplius: a larval stage in the life cycle of a
crustacean. Plural is nauplii, as in the larval,
just-hatched stage of Artemia spp. or brine
palytoxin: one of the deadliest natural toxins
known. In the aquarium, found in some en-
crusting anemones, especially in the genera
Palythoa and Protopalythoa. (Less commonly
in the genus Zoanthus.) All zoanthid-type
polyps should be handled with care, using
rubber or surgical gloves.
pantopods: so-called sea spiders or pycno-
gonids that are not true spiders. Sometimes
appear in aquariums as hitchhikers on live
rock or with zoanthids and other cnidarians.
They lack the venomous bite found in some
true spiders.
sessile: in marine biology, an organism that is
fxed in one place and not free to move about,
such as a coral. The opposite of motile.
zeolites: naturally formed rocklike alumino-
silicate minerals used as adsorbents in many
processes. In the aquarium, they are used by
some to remove ammonia and other nitrog-
enous dissolved compounds.
Techni cal terms that
appear i n arti cl es i n thi s i ssue lexicon|
REEF LIFE page 130
Wakatobi Marine Park, Southeast Sulawesi, Indonesia
Nudibranch (Hypselodoris apolgema) photographed
from underneath as it deposits its egg ribbon while on
the ceiling of a cavern.
DENISE NIELSEN TACKETT, co-author, with Larry P.
Tackett, of REEF LIFE, Natural History and Behaviors of
Marine Fishes and Invertebrates (Microcosm/TFH, 2002).
CRF Staghorn Nursery
Key Largo, FL
Photo by Luke Popwell
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ishing is fundamental to the Hawaiian culture.
It has sustained native Hawaiians for genera-
tions, and today shing in Hawaii is a vital com-
mercial and recreational activity that offers sig-
nicant economic value to shers and a viable
model for other sheries around the world. Myriad in-
dustries benet from Hawaiis reef and ocean resources.
The marine aquarium sherythat shery in which ani-
mals are collected live for the marine aquarium tradeis
the states most valuable and most studied inshore sh-
ery. It is also one of the most controversial.
For some, the question at the heart of the debate
about Hawaiis aquarium shery is one of sustainability.
All types of shing harvest animals from the ecosystem.
Most are intended as food sh, although in food sher-
ies a disquieting number of shes and other sea life or-
ganisms are taken unintentionally as bycatchanimals
caught and lost to mortality in the pursuit of more com-
mercially valuable species.
Marine aquarium sheries worldwide take a tiny
fraction of the biomass harvested by food sheries, yet
marine aquarium sheries often attract a disproportion-
ate number of emotional attacks. In Hawaii, this criti-
cism has taken a nasty turn: it has become an intense
dispute involving lawmakers and lawyers, lobbyists, and
activists, and it could shut down the aquarium shery.
I started covering Hawaiis marine aquarium shery for
CORAL magazine in the fall of 2010. I have gone diving
with aquarium shers, sailed aboard dive vessels, spent
time in the eld with state sheries biologists, interviewed
lawmakers, had coffee with leading antiaquarium-trade
activists, talked story with the keepers of native island
traditions, and generally tried to develop the most com-
prehensive view of the aquarium shery possible.
I have come to believe that much of the opposition to
the aquarium trade in Hawaii is really about ethics, not
sustainability. While anti-trade activists often wrap their
attacks in the guise of concern about resource sustain-
ability, the truth is that theirs is a very human and very
emotional opposition.

advanced aquatics | RET TALBOT
High drama for the
Hawaiian aquarium trade
In large part, this emotional
argument against the shery has
emerged from the island of Ha-
waii (the Big Island), where
the majority of aquarium sh-
ing occurs, and is directly related
to that islands geography, with
its narrow fringe of relatively
shallow reefs close to shore.
This constricted band of near-
shore reef habitat tends to force
a greater degree of user conict
than one generally sees on the
farther-ung reefs of Oahu,
home of Hawaiis second largest
aquarium shery.
Usersbe they divers, shers,
boaters, or any other group de-
ned by the activity it performs
in near-shore watersusually
identify most closely with others
in their user groups. Divers, for
example, dive together, shop at
the same dive shops, interact on-
line in dive forums, and read the
same dive publications. The same
could be said for aquarists. Being
part of a group often leads to a
sense of community, and a group
or community is frequently de-
ned by its relationship to other
groups and communities. When
conicts arise, these camps can
become polarized, and that is ex-
actly what has happened on the
Big Island, where divers and aquarium shers often do
not get along.
It is morning on a beautiful bay along
the west coast of the Big Island. The
bay is so calm that one person, look-
ing over the side of the dive boat,
says, Its like looking into an aquar-
ium. A variety of individual species
can be picked out as they move in
and out of the reef structure. There
are pairs of butteryshes, lone par-
rotshes, and scores of Humuhu-
munukunukuapuaa (Rhinecanthus
rectangulus) and other triggershes.
While this kaleidoscope of reef life
is amazing, it is the ashing shoals
of Yellow Tangs that elicit the biggest
response from the divers as they suit
up and prepare to go over the side.
Nearby, an aquarium sher snugs his anchor rope
and gets ready to dive. He has shed here beforeit is
a legal site for aquarium shing within the West Ha-
waii Regional Fisheries Management Area (WHRFMA).
While the tourists on the dive boat enter the water with
cameras and little plastic sh ID cards, the sher goes
over the side with a barrier net, a hand net, and some
collection buckets. Having spent more days in a year div-
ing these reefs than the dive tourists will likely spend in
a lifetime, he does not need sh ID cards, for he knows
the species and their behaviors well. Most dive tourists
do not have the same familiarity; one told me that the
opportunity to dive Hawaii was a lifelong dream real-
ized. Most have traveled thousands of miles to get here;
the sher had a 20-minute commute from the dock at
Honokohau Harbor where he keeps his boat, a trip he
makes several days a week when the weather is good.
The divemaster leads the tourists over a dive plan
as familiar to him as the easy morning commute is to
the sher. All the usual marine suspects are presentso
much so that one of the tourists who has dived at this
site before jokes that the animals must be animatronic
puppetsLike at Disneyland, she laughs. The divers
marvel at the apparent charisma of many of the species
they encounter as they n along the edge of the reef crest.
The sher, an extraordinarily skilled diver, is aware of the
dive tour before they see him. The divemaster points out
the sher, and one of the tourists takes a picture of him,
the ash of the strobe punctuating the brief encounter.
The sher turns back to his net and the task at hand
collecting Yellow Tangs for the marine aquarium trade.
Back on the surface, one of the tourists asks what
the other diver was doing. Raping the reef, the dive-
master responds without hesitation. He then goes on to
give his take on the aquarium shery. His explanation
holds a lot of sway with the dive tourists, many of whom,
although sympathetic to what they have just heard, will


Hanauma Bay
Nature Preserve
on Hanauma Bay.
Conficts between
snorkelers, divers,
and fshermen
have spawned
fshing reform
Yellow Tangs, Hawaiis number one live species export.
eat mahi-mahi, tuna, snapper, and other locally caught sh for dinner that
night. They may even discuss how horrible the aquarium trade is while slicing
through their Yellown steaks. Most do not see any contradiction because
they are more familiar with tuna steaks than the fusiform shape of the liv-
ing animal. Besides, a tuna is not as charismatic as an Ornate Butterysh
(Chaetodon ornatissimus).
Robert Wintner, owner of Snorkel Bobs and at one point the de facto
leader of the antiaquarium-shery movement in Hawaii, once told me he
does eat Hawaiian shes like mahi-mahi. I asked him if he saw a contradic-
tion in opposing the aquarium shery while being a consumer of species
caught by the food sheries, where serious concerns exist regarding bycatch
and sheries management is questionable. He started to say noone is about
sustenance and the other is what he calls an amusementbut then he hesi-
tated and told me about a personal encounter he had had with a mahi-mahi
while diving that, he allowed, may change his eating habits.
Choosing not to eat shor any other animalbased on personal ethics is
something I can respect, so long as we are clear that such a choice is about
ethics. An important truism is a shery is a shery is a shery. Whether
it is a recreational speargun or a hook-and-line shery, a commercial food
shery or an aquarium shery, sheries remove shes from the ecosystem. To
maintain healthy populations, shers must not catch more shes than the
reef can produce.
Most shes reproduce in such a way that, given the incredibly fecund and
competitive world in which they live, there are normally plenty of surplus
animals to replace those individuals that do not survive. The shery targets
the surplus, and sustainable sheries management is all about ensuring that
there are a sufcient number of adults in the shery to produce enough off-
spring to withstand shing pressures. Its simple as that, but the devil is in the
detailsand when the user conict escalates, as it has steadily over the past
three years in Hawaii, everyone wants to argue about whose details are facts.
Separating the facts from the fantasies is not terribly difcult when it
comes to Hawaiis marine aquarium shery: it is the best-studied inshore
shery in the state. In fact, it is one of the states best-studied sheries, period.
We have a lot of data, says Dr. William Walsh, the Big Islandbased
state sheries biologist who has the most comprehensive experience with the
marine aquarium shery. Based on that data, we have a pretty good idea
whats going on out there. When I rst interviewed Walsh back in 2010, I
asked if he thought the shery was being shed sustainably. It can be, but
to get to that point sheries managers need more tools at their disposal,
he explained. Several rule changes in the WHRFMA, including a 40-species
White List of approved shes for collection, bag and slot limits on some spe-
cies, and a host of other management measures, are intended to ensure con-
tinued resource sustainability, enhance near-shore resources, and minimize
user conict.
I have heard a lot about these proposed new rules from all sides of the
argument, especially when I spoke in the spring of 2012 to the West Hawaii
Fisheries Council (WHFC), the multi-stakeholder group that facilitated the
rules package. By this point, I was getting worried that my audience might be
losing patience. The proposed rule changes, which had been over a decade in
the making, seemed to be forever on the verge of becoming something more
than a proposal. If I was growing weary of the wait, however, I could only
imagine what the state biologists and members of the WHFC were think-
ing. A decade of committee meetings, research, and community discussion
involving more than 550 community membersthats whats at stake, one
WHFC member said.
For anti-trade activists like Robert
Wintner (Sea Shepherd), Rene Um-
berger (For the Fishes), and a vocal
group of their supporters, who tes-
tify before county and state legislative
bodies in support of closing the aquar-
ium shery, there is a lot more at stake
than a multi-stakeholder process. They
argue that Hawaiis reefs themselves
are in danger, and on that count they
are righttheir concern for ecosystem
health is warranted. Hawaiis reefs are
suffering from a crush of afictions.
Some may be natural, but others are
the result of anthropogenic stressors
like development, terrestrial runoff,
and an increasing number of users all
staking claims to the same resource.
While Dr. Walsh and his colleagues
believe that changes are needed to bet-
ter manage the aquarium shery, he
says the data clearly does not point
the nger at the aquarium shery as
the primary culprit in Hawaiian reefs
This January, a whole bevy of bills
seeking to ban or regulate the aquar-
ium shery were introduced, many by
legislators who tell me they were sim-
ply listening to their constituents.
Unlike last year, several of these bills
quickly made it to committee hear-
ings, and shers and those in support
of the marine aquarium shery are
very concerned that the shifting tide
in the Hawaii State Legislature may
not be going their way.
At the time of this writing, the
legislators dealing with these bills ap-
pear to be listening to the sheries
managers and actually looking at the
data. Several bills seek to regulate the
shery more rigorously, and a few of
these have widespread support from
both shers and others who support
Hawaiis marine aquarium shery. The
proposed rules package will nally be
making its way to the Board of Land
and Natural Resources in the coming
months. While the ght over the rules
grabbed a lot of headlines, the testimo-
ny submitted shows very strong sup-
port for adopting the rules into law
A&M Aquatics . . . . . . . . . . . . . . . . . . . . . . . . . . . 15, 52
Acan Lighting . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .103
AlgaGen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
American Marine . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Aqua Craft Products . . . . . . . . . . . . 5, 17, 107, 126
Aqua Medic . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81, 117
Aquascapers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .123
Aquatic Pixels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .127
Atlantis Aquariums . . . . . . . . . . . . . . . . . . . . . . . . .123
Bashsea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .108
Boyd Enterprises . . . . . . . . . . . . . Inside front cover
Breeders Registry . . . . . . . . . . . . . . . . . . . . . . . . . .119
Brightwell Aquatics. . . . . . . . . . . . . . . . . . . . . . . . . . 21
Champion Lighting & Supply . . . . . . . . . . . . . . . . . 96
Continuum Aquatics . . . . . . . . . . . . . . . . . . . . . . . . . 85
Coral Magazine Sources . . . . . . . . . . . . . . . . 122, 123
Coral Magazine Subscriptions . . . . . . . . . . . . . . . . 18
Coral Restoration Foundation . . . . . . . . . . . . . . .122
CPR Aquatics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
D-D . . . . . . . . . . . . . . . . . . . . . . . . . . inside back cover
EcoTech Marine . . . . . . . . . . . . . . . . . . . . . . . . 10, 11
Fauna Marin/Reef Wholesale . . . . . . . . . . . . . . . . 23
Fluval Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Fritz Aquatics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .106
Grotech . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Hydor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Instant Ocean . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Karen Talbot Art . . . . . . . . . . . . . . . . . . . . . . . . . . . .127
Kent Marine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Lifegard Aquatics . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Lifereef Filter Systems . . . . . . . . . . . . . . . . . . . . . .104
MACNA 2013 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .119
Marata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .107
Marine Breeding Initiative . . . . . . . . . . . . . . . . . .119
Milwaukee Instruments . . . . . . . . . . . . . . . . . . . . . . 14
Ocean Nutrition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Ornamental Fish Health Symposium . . . . . . . . . . 98
Orphek . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Pet Advantage . . . . . . . . . . . . . . . . . . . . . . . . . . . . .123
Piscine Energetics . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Poly-Bio Marine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
PolypLab . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .114
Prodibio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .118
Quality Marine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Red Sea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Reef Aquaria Design . . . . . . . . . . . . . . . . . . . . . . . .100
ReefBuilders . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .107
Reef Nutrition . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Reefs.com . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Rising Tide Conservation . . . . . . . . . . . . . . . . . . .113
Rods Food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .114
San Francisco Bay Brand . . . . . . . . . . . . . . . . . . . . . 30
SEAMIX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22, 129
Segrest Farms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .105
Stax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Thrive Aquatics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Tropic Marin . . . . . . . . . . . . . . . . . . . . . . . . back cover
Tunze . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Two Little Fishies . . . . . . . 16, 26, 29, 57, 98, 108, 114
Ushio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Wallet Pen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .119
ZooMed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

For a CORAL Media Kit or other information, please contact:
[amcs Lawrcncc, Fublislcr - 802.985.9977 Ext. 7 - [amcs.Lawrcncc@Rcc2Rainorcst.com
which means that aquarists should go back to keeping their aquaria and not
worry so much about the battle over Hawaiis aquarium shery, right? Wrong.
Hawaiis marine aquarium shery is a model shery in which managers
are managing the stocks based on data and where shers, and others who
use and care about the reefs, are attempting to address user conict through
a multi-stakeholder process. It is not perfect, but aquarists must understand
how different this shery is from most of the Indo-Pacic marine aquarium
sheries that provide aquarium animals to sh stores across the country.
Hawaiis aquarium shery is, for the most part, operated sustainably. It is
a shery in which the use of destructive shing practices is rare. It is a shery
whose shers are generally appropriately compensated for their efforts. It is a
shery that highly values its animals, resulting in better care from the point
of collection to the point of sale in the local aquarium shop.
Unfortunately, the majority of marine aquarium sheries in places like
the Philippines and Indonesia are not managed based on data; many are not
managed at all in any real sense of the word. They are frequently locally over-
shed, and destructive shing practices like using cyanide are still employed.
Fishers in these countries commonly do not earn a fair wage, and the animals
are often devalued along the chain of custody, resulting too frequently in
compromised animal health and unnecessary mortality.
So when we aquarists think about Hawaii, we should also think about
what type of aquarium shery we wish to support with our purchasing power.
Understanding what makes Hawaiis marine aquarium shery a model sh-
ery can lead to immediate actions that will initiate real change in the trade.
While exporting Hawaiis model in its entirety is probably not realistic at
present, aquarists can do their part to support and encourage sustainable
marine aquarium sheries throughout the developing world, in places like
the Solomon Islands, Fiji, and Papua New Guinea. Supporting these sheries
can create tangible economic incentives for shing sustainably and conserv-
ing reef ecosystems and promote positive and lasting socioeconomic benets
to shers and their communities. It can make a difference, but it must begin
with us.
Ret Talbot is a CORAL senior editor who writes frequently about sustainability issues.
He lives in Rockland, Maine with his wife, conservation artist Karen Talbot. J. Charles
Delbeek will return to this space in the next issue of CORAL.


Tikis Honaunau
Place of Refuge,
Big Island. Hawaii
is a place of
ancient fshing
traditions and
The ultimate high
magnesium salt
evaporation of water taken from one of the richest coral
seas on the planet. This results in a salt in which every
bucket contains over 70 trace elements in exact natural
proportions including 23 which occur at less than 1 PPM.
This pure base salt is then specially enhanced for the reef
aquarium by the elevation of specic parameters required
for growth and colour such as magnesium, calcium,
potassium and dKH. The result is a unique formulation
which gives you fantastic results.
Even if you can detect all of the elements that occur naturally in the water
around the reef and determine the levels correctly, imagine attempting to blend
these 23+ minor trace elements evenly during the manufacture of a synthetic
salt when they occur at less than 1 gram to 1 tonne of salt. What is the effect of
these trace elements if you get more than your fair share in your bucket? With
H2Ocean Pro+ we let nature be your mixing pot so we guarantee you every
bucket is correct.
GUARANTEED PARAMETERS (salinity 35. 5ppt)
The formulation for H2Ocean Pro+ salt was developed to give you the optimum
chemistry for a healthy reef aquarium and to allow growth of even the most
difcult corals and sponges and to date the demand for this salt and the
positive feedback from both hobbyists and experts alike have exceeded our
Many salts concentrate on enhancing calcium levels and often ignore the
importance of magnesium. The correct magnesium level has an enormous
impact on how easy it is to maintain the calcium level, pH and alkalinity in your
tank and can halve the time that you need to run your calcium reactor.
We recommend that you use a D-D portable SEAWATER refractometer for
accurate and consistent measurement of the salinity in your aquarium. These
are widely available from your D-D retailer.
H2Ocean Pro+ is designed for use with reverse osmosis, deionised or soft
water with a calcium level below 30mg/l. As the calcium level in H2Ocean Pro+
is already boosted to 440mg/l then mixing it with hard tap water containing
additional calcium may exceed the point at which it will precipitate out
of solution.
Reverse osmosis removes ions such as nitrate and phosphate from your
tap water which otherwise would contribute towards nuisance algae in
your aquarium.
8.2 - 8.4
8.7 - 9.8
Calcium (Ca2+)
430 - 460
Magnesium (Mg2+)
1300 - 1380
Chloride (Cl-)
19960 - 20130
Potassium (K+)
380 - 420
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