unclenedsfishfactory.com

LETTER
Long-term trends of coral imports into the United States indicate
future opportunities for ecosystem and societal benefits
Andrew L. Rhyne1,2, Michael F. Tlusty1, & Les Kaufman1,3,4
1 New England Aquarium, John H. Prescott Marine Laboratory, Boston, MA, USA
2 Roger Williams University, Department of Biology and Marine Biology, Bristol, RI, USA
3 Boston University Marine Program, Department of Biology, Boston University, Boston, MA, USA
4 Conservation International, Arlington, VA, USA
Keywords
Aquarium trade; coral trade; curio trade; coral
triangle; marine policy.
Correspondence
Andrew L Rhyne, Department of Biology and
Marine Biology, Roger Williams University, One
Old Ferry Road, Bristol, RI 02809, USA. Tel: 401
254-5750; Fax: 401 254-3310. E-mail:
arhyne@rwu.edu
Received
5 March 2012
Accepted
12 June 2012
Editor
Dirk Roux
doi: 10.1111/j.1755-263X.2012.00265.x
Abstract
The international trade in corals used to be primarily a curio trade of dried
skeletons, but now focuses on live corals for the marine reef aquarium trade.
The trade is still rapidly evolving, creating challenges including the addition of
new species that outpace effective management strategies. New species in the
live coral trade initially command high prices, but as they become common the
price radically decreases with feedback effects to the trade. To understand these
trends, 21 years of live coral import data for the United States were assessed.
Trade increased over 8% per year between 1990 until the mid-2000s, and has
since decreased by 9% annually. The timing of the peak and decline varies
among species, and is a result of the rising popularity of mini-reef ecosystem
aquariums, the global financial crisis, and an increase in aquaculture production.
The live coral trade offers opportunities for coral reef ecosystem conservation
and sustainable economic benefits to coastal communities, but realization
of these externalities will require effective data tracking.

Introduction
The marine aquarium trade is but one of many threats
facing the world’s coral reefs, and despite some public
opposition to the trade (reviewed in Tlusty et al. 2012),
few quantitative studies of its true significance have been
reported (Kolm & Berglund 2003). Species entering the
market are extremely diverse and include algae, vascular
plants, a wide range of invertebrate phyla and both bony
and cartilaginous fishes. The trade has the potential to
cause overharvesting, collateral damage to coral reef
habitat and introduction of exotic species (Smith et al.
2008; Jones et al. 2009; Rhyne et al. 2009; Schofield et al.
2009). However, it is also a potential boon to conservation
through elevated valuation and stewardship of
wild habitats that support livelihoods based upon the
collection, rearing and export of aquarium specimens
(Tlusty 2002; Tlusty et al. 2008). The aquarium hobby can
also benefit conservation by mobilizing public pressure
to rein in other anthropogenic stressors including global
climate change (Carpenter et al. 2008), unsustainable
global market demand (e.g., shark fin, Napoleon wrasse,
Chinese traditional medicine), and development of
coastal and aquatic resources. Combined with commercial
overfishing (Burke et al. 2011), these nonaquariumrelated
threats are likely to be far more severe and pervasive
than the direct damage done by aquarium collectors,
though comparative data are lacking. Consequently, we
have launched a systematic study of the marine aquarium
trade and its current and potential relationship to
coral reef conservation. Here we report on the changing
face of the trade, in nature and magnitude, over the past
21 years with particular regard to coral biodiversity.
The export of marine aquarium specimens is a global
industry (Wabnitz et al. 2003). Each year, a complex trade
network sells upwards of 50 million coral reef animals
to an estimated 2 million hobbyists worldwide (Wood
2001; Green 2003; Wabnitz et al. 2003). The epicenter of
478 Conservation Letters 5 (2012) 478–485 Copyright and Photocopying: c 2012 Wiley Periodicals, Inc.
A.L. Rhyne et al. Trade of aquarium and curio corals
trade is the Coral Triangle, a region of high biodiversity
that encompasses Indonesia, Malaysia, Papua New
Guinea, Philippines, Solomon Islands, and Timor-Leste.

The Philippines and Indonesia represent about 85% of
trade volume, however, smaller island nations export
unique species helping to drive high rates of overall trade
biodiversity (Rhyne et al. 2012). Over 40 countries export
livestock to feed the hobby and for the past three decades
growth in traded volume exceeded 10% per annum
(Bruckner 2001), with some slowing at the end of 2007
as a result of the global economic recession (Rhyne et al.
2009). The United States is a major destination of live
coral reef products importing 68% of all live corals reported
in CITES with the EU importing 24% (mostly split
among Germany, France, the Netherlands, and Great
Britain) (Jones 2008). Given the significant U.S. role in
the coral trade, the United States Coral Reef Task Force
identified reforms to the marine aquarium trade as a major
priority in 2000 (www.coralreef.gov/international/).
This resulted in efforts to increase the knowledge of this
trade within the United States (Puerto Rico and Hawaii)
and to help promote a sustainable global trade internationally
(Tissot and Hallacher 2003; Legore et al. 2005).
While there are many management challenges for
ornamental fisheries at the local and the national level,
global trade is the key driver (Green 2003; Olivier 2008).
This is most evident in the coral trade (Bruckner 2001;
Moore and Best 2001; Tissot et al. 2010), which began
many decades ago with the harvest of dried “curio”
specimens. The large take resulted in management measures
focused on the total mass of harvested coral (Lovell
2001). With the advent of the marine home aquarium,
trade emphasis shifted to the collection and transport
of a much smaller volume of live colonies from remote
regions (Figure 1). During the mid-1980s, scientists
began to note a rapid, global degradation of coral reef
habitats, an event in which the curio and aquarium trade
were clearly participants. Because of the awareness of
impacts to corals, “stony corals” and “live rock” have
been protected by the Convention on International trade
in Endangered Species of Wild Fauna and Flora (CITES
code AC22 Inf. 8) since 1985, and regulated by export
quotas and stringent import regulations (Department of
the Interior Fish & Wildlife Service 2007). “Soft corals”
and attached coral “substrata” are not covered by this
convention and therefore are not highly regulated. The
degradation of reefs and concern about trade of corals
led to the ban of shipments from the Philippines after
the CITES listing (reviewed in Green & Shirley 1999).
Since the CITES listing, the harvest of corals and coral
rock has seen a dramatic shift; moving away from a
curio (dead) based industry to the trade of live coral for
the marine reef aquarium trade. Indonesia, the largest
1980-1995
Aquarium trade
-wild fishery
-remote locales
-increasing volumes
- ornamental value
Post-1995
Aquarium trade
-rare species
-wild fishery
-remote locales
-decreasing
-common species
-aquaculture
-central production
-increasing
?
Future
-aquaculture of curios
-MPA protection of
slow growing species
-aquaculture production
for habitat enhancement
-economic benefits for
remote locales
Pre 1980
Curio trade
-wild fishery
-remote locales
-large volumes
-little value for rarity
Figure 1 The trade in corals for the curio and live home hobby trade. The
curio trade was the most important until the marine home hobby trade
became popular. With the advent of ecosystem reef tanks in the mid-
1990s the trade changed again, with aquaculture of fast growing corals
presently increasing. Will the future allow for aquaculture production of
fast growing corals to supply the curio trade? How can the coral fisheries
be managed to provide economic benefits to the remote communities
dependent originally upon the fishery?
supplier of stony corals, along with other south Pacific
countries, claimed harvests did not significantly affect
wild populations, and implemented harvest quota systems
in an effort to comply with CITES (Harriott 2003).
While quotas provide an important trade limitation for
threatened species (Harriott 2003), questions on their
effectiveness have been raised (Bentley 1998; Castello &
Stewart 2010; Knittweis & Wolff 2010), and the initial
large quotas may not be sufficient for slow growing stony
coral species (Bruckner 2000). Bruckner (2001) reviewed
the strengths and weakness of CITES for monitoring the
international trade of marine aquarium species, citing the
key limitation as the lack of taxonomic level information
(Green & Hendry 1999). The major concern is that rare
species could be overexploited under quotas set at the
genus level. The illegal trade of corals is difficult to track,
as it is for all wildlife, and hence significant portions of
the trade enterprise can potentially be unaccounted for
in formal data (Rosen & Smith 2010; Toledo et al. 2012).
However, there will be an asymmetry in the illegal trade
of corals. More illegal trade will likely occur in the curio
side as the handling of live coral through known shipping
routes makes it more difficult to hide this trade compared
to the bulk cargo shipping of curio products.
The aquarium trade is a rapidly evolving industry
(Larkin 2008; Olivier 2008; Rhyne et al. 2009). Species
Conservation Letters 5 (2012) 478–485 Copyright and Photocopying: c 2012 Wiley Periodicals, Inc. 479
Trade of aquarium and curio corals A.L. Rhyne et al.
can enter the trade and quickly become heavily traded
(e.g., Banggai Cardinalfish, Pterapogon kauderni) (Lunn &
Moreau 2004). Also, species that have been traded at a
low level for decades can suddenly become a new fad and
experience a sharp increase in exploitation (e.g., Ricordea
mushrooms, Ricordea florida) (Torres-Pratts et al. 2011).
As new products move into the market, retail prices
are initially high until supply increases, at which point
prices dramatically drop (Larkin 2008). Because of the
low price, there is typically an influx of low quality supply
as the species becomes popular and more consumers
can afford the cheaper product. This rapidly changing nature
of the aquarium trade creates numerous management
challenges relevant to the trade of live coral. For
example, the rapid shift in markets might allow for the
over exploitation of slow growing corals harvested under
management systems designed for faster growing corals
(Jones 2011). Here, we analyze trade data for live corals
for two decades from 1990 to 2010. The purpose of this is
to understand the overall trends over time as coral species
enter the trade and become commercially popular. We
also assess the impact increased aquaculture production
of corals (Parks et al. 2008) has on the trade, and forecast
how changes in the trade of live corals may impact coral
reef ecosystems.
Methods
While the focus of the aquarium trade has historically
been on fishes, the advent of live reef and mini-reef
aquarium set-ups has greatly increased demand for
marine invertebrate taxa (Rhyne et al. 2009). Arthopoda,
Mollusca, and Cnidaria are the most heavily traded tropical
marine invertebrates (Wabnitz et al. 2003). The most
popular cnidarians (approximately 5% of the total trade)
are corals and anemones with a few species of jellyfishes
(Wabnitz et al. 2003). Corals within the aquarium trade
can be loosely defined as polyps (corallimorphs, and
zoanthids), gorgonians, and both soft and hard corals.
They are categorized arbitrarily based upon the presence
or absence of a hard skeleton rather than on the
basis of actual phylogenetic relationships. For example,
species of Alcyonacea, Corallimorpharia, Gorgonacea,
and Zoantharia are often lumped into the general term
“soft corals,” while the anthozoan orders Scleractinia and
Helioporacea and hydrozoan order Anthoathecata are
traded as “hard corals.” In the same vein, some species of
the Alcyonacea, namely Tubipora spp., are traded as hard
corals.
To better understand the recent changes in the stony
coral trade and assess market trends we queried CITES
trade data for a 21-year period (1990–2010) utilizing the
World Conservation Monitoring Center trade database
(United Nations Environment Programme–World Conservation
Monitoring Centre 2010). Because the United
States is the world’s largest importer of stony corals
(Jones 2008) and numerous authors have noted discrepancies
between import and export data in CITES
(Bruckner 2001; Blundell & Mascia 2005; Jones 2008;
Phelps et al. 2010), we limited our queries to imports
of corals reported by the United States to CITES for
all countries. To compare the long-term trends in the
commercial curio and live trade of stony corals we analyzed
“purpose code” trade data for trends in both “live”
and raw or worked corals. We limited the possibility
of taxonomic misidentification by sorting all records to
genus level and only analyzed the aquarium corals in the
reports that listed “live” items and not kilograms. Live
corals traded in kilograms are likely to be biologically
active “live rock,” and not coral species of interest to the
hobbyists. For example, the more than 300,000 kg of
“live” Acropora exported from Tonga in 2003 is almost
certainly a live rock product sold under the terms “Tonga
branch” or “shelf rock.” However, it is not live Acropora,
it is likely dead, encrusted fragments of branching coral
skeleton. The CITES data were used to designate corals
by common trade designations (small polyp, SPS, or large
polyp, LPS), life history traits (free-living or stationary)
and method of production (wild or cultured) as indicated
by CITES source codes. In an effort to understand how
retail price was related to the volume traded, price data
were gathered for all genera imported into the United
States by collection price data off the internet available on
retailer web pages as well as eBay. Listing on eBay was of
specific interest because of the number of rare or unique
corals being offered for sale. Retail prices (price per individual)
were analyzed against trade volume, life history
traits (free, fixed) and trade designations (SPS, LPS) and
variation (more than 10%) from mean retail prices were
further scrutinized for likely causes for this deviation.
Results and discussion
Trade
The total number of stony corals imported into the United
States from the Coral Triangle and South Pacific grew at a
yearly rate of 8.75% from 1990 until the mid-2000s. After
a brief level period it dramatically declined by 8.98%
annually (Figure 2). During this period, the proportion of
corals originating from the Coral Triangle decreased from
>90% in the 1990s to a low of 65% in 2010 as Australia
expanded its exports. The life history traits (free, fixed)
and trade designations (SPS, LPS) of coral species imported
into the United States can be combined into three
480 Conservation Letters 5 (2012) 478–485 Copyright and Photocopying: c 2012 Wiley Periodicals, Inc.
A.L. Rhyne et al. Trade of aquarium and curio corals
Figure 2 Number (individual items unless otherwise indicated) of corals
imported into the United States over the past 21 years by trade type (top)
for live species grouping (middle) and by% composition (bottom). LPS and
SPS are trade terms for Large Polyp Stony corals and Small Polyp Stony
corals. LPS species are grouped into Fixed, those species that attached
to substrata or Free those species that are free living and not attached
to the substrata. Raw and worked (curio trade) corals (top) are shown in
both kilograms and pieces. Note the large increase in kilograms imported
in 2003 is a result of live rock (Tonga Branch Rock) imports mislabeled as
raw Acropora sp. from Tonga.
groups (Figure 2), each exhibiting a different historical
trend. The percent of SPS-fixed corals in the trade tracks
the total import numbers, increasing from <10% in 1992
up to 38% in 2006, then decreasing to 27% in 2009. The
LPS-fixed corals trend in the opposite direction, decreasing
between 1992 (>50% of the trade) and 2006 (34%)
before rising again to 43% in 2009. The percent of LPSfree
corals in the trade decreased between 1992 and 2009
beginning at just over 38% and ending at just under 30%
of the trade (Figure 2).
This pattern is repeated in the annualized import
records of the top six species of stony corals (Figure 3).
Acropora spp. imports decline sharply post 2006 while the
Figure 3 Top six genera of stony corals imported as “live,” in individual
units, for trade into the United States over the past 21 years.
remaining five species (all SPS) peak earlier and decrease
slightly if at all. The increase in the number of species
imported is a result of the switch in the marine aquarium
hobby to the keeping of “mini-reef” ecosystems (Rhyne
et al. 2009). This trend in the hobby also explains the
change in the proportion of the different types (positive
trend of SPS and negative of LPS), as SPS are more desirable
for the smaller mini-reef aquariums. The dramatic
decrease post-2006 was likely a result of two factors. The
first was the economic downturn (Mittelman 2009), in
which spending on “luxury items” was curtailed. Probably
more important, however, was a change in coral
sourcing. While the majority of corals are collected from
the wild, there is a small but growing fraction of corals
from aquaculture production (Figure 4). Nearly all-wild
corals imported into the United States for the aquarium
trade originate from Pacific sources, with a majority
originating from the Coral Triangle, although Fiji and
Australia are shipping greater proportions of the total
amount. Currently, only a small number of coral genera
can be commercially produced in aquaculture with a
majority belonging to the genus Acropora (Figure 4b). The
significant decline in Acropora spp. imports to the United
States (Figures 3 and 4b) is a result of the switch in
production from wild capture to aquaculture. Hidden in
this decline is a growing importance of ex situ production
through fragmentation (“fragging”), which has been
successfully used for commercial production of SPS and
some branching LPS corals. Much of this type of production
can occur within the United States and import
records do not account for domestic production, which
began in the early 1990s (Perrin 1993). Currently, the
statistical tracking of coral production inside the United
States is lacking. Thus, the U.S. domestic production
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Trade of aquarium and curio corals A.L. Rhyne et al.
Figure 4 (Top) Total number of individual corals imported into the United
States. (Bottom) Acropora sp. imported into the United States. White bars
are from wild harvests,while black bars indicate those corals from captive
culture (CITES source codes F, R, C).
of SPS corals is a likely factor in causing the negative
trend in both the percent imports of SPS-fixed species, as
well as the total number of corals imported. Increasing
aquaculture production also occurs in source nations. For
example, 26% of Acropora imported into to the United
States from Indonesia in 2009 were listed under CITES
codes (F-farm, C-captive bred, and R-ranch) indicating
aquaculture production.
Aquaculture will likely continue to gain traction with
the vast majority of SPS corals being aquacultured within
the next decade. Not all species lend themselves to culture,
however. Large polyp species of corals in the sessile
genera Catalaphyllia, Nemenzophyllia and all free living
LPS corals are more challenging to produce and therefore
wild production is likely to dictate supply for the time
being. Free living species, which can be fragmented commercially,
e.g., Fungia spp. are difficult to certify as aquacultured
according to existing standards (Pasaribu 2008)
and cultured species may be imported under existing wild
harvest quotas.
% Australian
Figure 5 (Top) Relationship of price and volume (total number imported)
for corals imported into the United States. (Bottom) Relationship (r2 =
0.55) of mean price and composition of coral imported from Australia.
Markets
Retail prices of all corals averaged US$56.29 per piece
ranging from US$5.00 to US$499.99. No relationship
was observed (y = –9E−05x + 57.56, r2 < 0.01) between
volume imported and average price (Figure 5 top). However,
SPS-fixed corals were more likely to be devalued
than the two LPS designations, as the residuals from the
above mentioned relationship for the SPS corals were
significantly less than 0, and lower than the residuals for
the LPS corals (Figure 6). The Australian corals had a
major influence on this relationship as the more prevalent
these corals became, the greater the price (Figure 5
bottom). The higher-priced corals are sold with key
words of “rare,” “limited edition,” “unique” or sold under
producer’s name or strain name (e.g., Green Warpaint
482 Conservation Letters 5 (2012) 478–485 Copyright and Photocopying: c 2012 Wiley Periodicals, Inc.
A.L. Rhyne et al. Trade of aquarium and curio corals
Figure 6 The average price range (US$) and the average residual from
the price-volume (total number traded) relationship (Figure 4 top) for the
three different coral categories.
Scolymia) which appears to have increased their mean
price due to limited number of highly priced individuals.
Therefore, prices appeared more dependent on perceived
market abundance and not the actual supply as indicated
from CITES records. While true abundance will affect
price (Courchamp et al. 2006), here it is postulated that
perceived rarity will also affect price, as often unusual
colors command greater prices than common colors. This
rarity price premium will also muddle a comparison of
the cost of wild and aquacultured species. Corals that
are sold as “aquacultured” are often much smaller than
wild corals with many domestic producers selling “frags”
or small cuttings of corals whereas wild corals are often
sold as “large” “x-large” or “show” sizes. Currently many
cultured corals are imported as wild product, since it is
simpler to identify aquaculture products as wild than it
is to provide all of the appropriate documentation of the
aquacultured product (D. Palmer, personal communication).
These confounding variables will make it difficult
to accurately determine the resource economics of the
coral trade.
Sustainability considerations
While these data are used to assess trade chains and the
value of the home hobby industry, they can also be used
to highlight where conservation programs can focus their
attention for maximum impact. The surge of aquaculture
amongst SPS corals indicates that these fast growing
corals would be ideal candidates for localized habitat
restoration programs (Edwards & Gomez 2007), or
even curio products. Typically, aquaculture concentrates
production near central distribution gateways, and not in
remote villages. Efforts need to be made to understand
the impact of this centralized production on small rural
communities in exporting countries, and to promote the
small communities that have been the historically important
harvest locales. This localized production along with
restocking would help to maintain coral reef ecosystem
services. Ornamental fishery products from many Pacific
island countries are a major source of export revenue,
and are important economic drivers, but it is important
to continue the financial benefits of the trade to the small
collector communities. Fijian communities that harvest
corals for the aquarium trade have a higher median
income from this harvest when compared to traditional
fishery products (Wabnitz et al. 2003; Lovell & Whippy-
Morris 2009). In situ aquaculture production or ranching
is a means to match the various needs for ecological, economic
and societal benefits. It is critical to have economic
benefits of ornamental species production (fishery or
aquaculture) remain in the original source communities
(Tlusty 2002): this should be a priority for a responsible
global trade, and will align the aquarium fishery with the
Convention on Biological Diversity (Secretariat of the
Convention on Biological Diversity 2005).
The CITES system for overseeing the live coral trade,
while imperfect as currently implemented, is a good one
and it accomplishes a majority of the objectives for trade
regulation (Phelps et al. 2010). Indonesia has responded
to CITES and calls for a more sustainable coral trade
by developing meaningful mariculture standards with
traceability (Pasaribu 2008). However, in a dramatically
fast moving industry, monitoring trends in the trade
is not a simple matter of assessing volumes over time.
For example, quotas that were derived from the curio
trade data in which product is measured in tonnage
and focused on fast growing species is inappropriate for
slower growing aquarium species that are sold by the
piece or “individual” (Jones 2011). Furthermore, price is
susceptible to product rarity and perceived supply limitations.
These “rare” corals do not appear in trade data, as
they are unique color morphs untraceable through the
CITES or any other governmental monitoring system.
The additional tracking of the value of imports will also
be important to determine if decreasing trends reflect
supply difficulties, which could indicate significant pressure
on populations from the fishery. If imports decrease,
and price remains stable or declines, then demand is
likely being met by domestic production. However, if
imports decline, and the price increases, then supply is
not meeting the demand (Courchamp et al. 2006).
Conclusion
In summary, the aquarium and curio trade for corals is
associated with positive and negative impacts to ecosystems
and societies. It is a rapidly evolving trade, and
because of this, needs to be adaptively managed in a way
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Trade of aquarium and curio corals A.L. Rhyne et al.
to support conservation efforts and maximize the benefit
of the trade. The increasing importance of aquaculture
will need to be integrated into both CITES rules and
also sourcing decisions for retailers within the trade.
Valuing in situ production of corals will help maintain
historically critical locales and communities, which will
in return maintain value to extant reef ecosystems. The
aquarium trade needs to be a fully vested partner in such
market-driven coral reef conservation initiatives.
Future monitoring must accurately catalogue wild and
aquacultured corals, their color morphs, and their value.
It will also be important to address (1) the relative risks
and benefits of ex situ or concentrated in situ production
of corals for the aquarium trade; (2) how significant
production increases will affect total revenue; and
(3) whether there can be a return to curio corals through
aquacultured eco- or fair-trade labeled products. Answers
to these questions will be critical to evaluate whether
coral production will continue to be part of a long-term
sustainable economic solution especially for remote island
communities.
Acknowledgments
The data and presentation of these data were prepared
for the 2011 United States Department of Commerce,
National Oceanic and Atmospheric Administration Coral
Mariculture Workshop held in Bali, Indonesia. Rhyne
is supported by a National Oceanic and Atmospheric
Administration (NOAA) Coral Reef Conservation Program
Grant NA09NMF4000385. The authors are grateful
to Ana Malone Oliver and Elizabeth Futoma, Roger
Williams University, for help in capturing the market and
CITES data. Laura Dee, University of California at Santa
Barbara, provided helpful comments on the market
data. Kerri Warren, Roger Williams University, and
two anonymous reviewers provide helpful comments
on a previous draft of this manuscript. This paper is a
contribution from the “Blueing the Aquarium Trade” initiative
of the New England Aquarium and Conservation
International.
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