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Douglas Fenner ÆKenneth Banks
Orange Cup Coral
Tubastraea coccinea
invades Florida
and the Flower Garden Banks, Northwestern Gulf of Mexico
Received: 13 May 2003 / Accepted: 3 January 2004 / Published online: 16 September 2004
ÓSpringer-Verlag 2004
Keywords Invasion ÆFlorida ÆCurrents ÆTubastraea Æ
Wreck ÆDiadema ÆFlower Garden Banks
Introduction
The azooxanthellate scleractinian coral Tubastraea coc-
cinea has recently been reported to have invaded the
Gulf of Mexico (Fenner 2001) and Brazil (Castro and
Pires 2001; Ferreira 2003; Figueira de Paula and Creed
2004). It may have been spreading in the Caribbean
since it was reported there by Vaughan and Wells (1943),
based on material from Curac¸ ao and Puerto Rico, and
may even have been introduced into the Caribbean from
the Pacific (Cairns 2000; Humann and DeLoach 2002,
p. 164). No Caribbean fossils of this species are known
(Cairns 1999). Most Caribbean reef building corals
(88%) were described before T. coccinea was reported
from the Caribbean in 1943, with the median date of
description being 116 years earlier. All other genera had
been found in the Caribbean before Tubastraea, the last
having been found 75 years before Tubastraea was
found. The type locality of T. coccinea is Bora Bora,
date 1829. Its range includes most or all of the tropical
Indo-Pacific (Cairns 2000). Alternatively, it may have
come from the Cape Verde Islands or Gulf of Guinea in
the eastern Atlantic, where it is also known (Laborel
1974; Cairns 2000). It has not been reported from
Florida (Wheaton and Jaap 1988; Jaap and Hallock
1990; Cairns 2000; Fenner 2001; Dawson 2002; see also
http://floridakeys.nos.noaa.gov/sanctuary_resources/
specieslist.pdf), and in the Gulf of Mexico it has only
been reported from oil platforms (Fenner 2001). It is
reported here for the first time in Florida and the Flower
Garden Banks of the northwestern Gulf of Mexico.
Results and discussion
T. coccinea was first observed on October 20, 2001 by
one of us (KB) on the steel tugboat ‘‘Donal G. McAll-
ister,’’ which was placed as an artificial reef at 21-m
depth about 2.2 km offshore from Hollywood, Florida.
A single orange-red colony, about 6-cm diameter was
observed on April 18, 2002, growing on the outside of
the cabin facing south at 16-m depth. Subsequently,
numerous colonies were discovered on 12 other sites in
southern Florida: nine other ships, one set of oil rig
jackets, a floating dock, and a limestone boulder reef
(Table 1). Many colonies were on the hulls of the ships,
in shaded areas. The coral was abundant on all four
wrecks that were examined for abundance: RSB-1,
Tenneco, Jay Scutti, and Capt. Dan (colonies were not
counted). The only non-shipwreck or dock sighting of
T. coccinea was a single colony on limestone boulders
(Port of Miami Mitigation Reef, Tim Mcintosh, per-
sonal communication). These boulders came from
upland quarries so the coral colony must have recruited
recently. The largest colonies observed on the artificial
reefs were about 6-cm diameter. A colony that was
collected was identified by Steven Cairns as T. coccinea.
The date at which T. coccinea first settled on these
structures has not been determined. Although growth
rates have not been reported in this species, aquarium
observations suggest they may reach a diameter of 5 cm
in about one year, and growth may slow in adults as
energy is put into reproduction with maximum sizes
reached of about 10–15-cm diameter (Julian Sprung and
Daniela Stettler, personal communication). The oil rig
Communicated by: Ecological Editor P. F. Sale
D. Fenner (&)
Australian Institute of Marine Sciences,
Townsville, Queensland, Australia
E-mail: dfenner@blueskynet.as
Tel.: +1-684-6334456
Present address: Department of Marine and Wildlife Resources,
PO Box 3730, Pago Pago, AS 96799, USA
K. Banks
Broward County Dept. of Planning and Environmental Protection,
218 SW 1st Ave, Fort Lauderdale, Fl 33301, USA
Coral Reefs (2004) 23: 505–507
DOI 10.1007/s00338-004-0422-x
jackets were transported to Florida on barges, making
the survival of any T. coccinea previously living on them
unlikely though perhaps not impossible (if parts were
immersed in water in the barge hull). Colonies on the
hulls of ships could have arrived with the ships.
Surprisingly, no colonies have been found on the first
artificial reef, the Mercedes, sunk in March 1985.
T. coccinea was first observed on the wreck of the
Duane off Key Largo by J. Sprung about 1999. Colo-
nies’ Were already numerous on it at that time. Colonies
about 5–10 cm diameter are now common on vertical
surfaces. This 100-m long steel ship’ W’ Which was sunk
in 1987, sits in water about 36-m deep with its top deck
at about 30-m depth. The Florida Keys National Marine
Sanctuary (FKNMS) website page (http://florida-
keys.nos.noaa.gov/sanctuary_resources/shipwreck_trail/
duane.html), last updated January 1, 2000, lists ‘‘Cup
Coral’’ (T. coccinea) as being on the Duane. The
FKNMS website also reports Cup Coral on another
wreck, the Amesbury (http://floridakeys.nos.noaa.gov/
sanctuary_resources/shipwreck_trail/amesbury.html,
last updated January 1, 2000). The Amesbury is located
8 km west of Key West in 8 m of water. J. Sprung also
observed one colony of about the same size on the
underside of a floating dock next to the seawall inside
Palm Beach Inlet.
T. coccinea was observed on the East Flower Garden
Bank of the northwestern Gulf of Mexico by H. Ly-
dersen-Bulman on August 28, 2002. A single colony
about 15-cm diameter was photographed at about 26-m
depth. T. coccinea was also observed on natural reef
ledges at about 10-m depth at Fowl Cay Preserve
(26°38.229 N 77°02.310 W) north of Man of War Cay,
Abaco, Bahamas, by KB in August of 2000, and abun-
dant colonies were found in caverns off the north end of
Guana Cay, Abaco (26°42.22 N 77°09.17 W) in 2003.
T. coccinea has now been reported to have invaded
the Gulf of Mexico, Brazil, and Florida. This is consis-
tent with the proposal that it has been expanding its
range in the Caribbean, and was originally introduced
into the Caribbean from the Indo-Pacific (Cairns 2000)
or perhaps the eastern Atlantic. It has been observed on
boat hulls in the Caribbean, where transport on boats
was suggested as a mode of dispersal (Cairns 2000), and
observed on ships in Brazil (Ferreira 2003, Ferreira et al.
2004). The dispersal route into the Gulf of Mexico was
not identified, though most likely it reached airplane
wrecks in the Caribbean by larval dispersal (Fenner
2001). It was first seen in the southeastern Gulf of
Mexico in 1977, western Gulf in 1985, Texas in the
northwestern Gulf in 1991, and Louisiana in the
northern Gulf in 1994, so larvae may have drifted with
currents clockwise along the continental shelf in the Gulf
of Mexico (Fig. 1). The relatively recent invasion of
Florida by T. coccinea could have occurred by transport
on the hulls of one or more of the ship wrecks it now
grows on, or it could have arrived as larvae on currents
from the Gulf of Mexico or Caribbean. The maps of
Roberts (1997) show that larvae could have reached
Florida from western Cuba in 1 month, and from
Cozumel in 2 months. Dispersal to Brazil was by mobile
oil drilling platforms (Castro and Pires 2001; Ferreira
2003; Figueira de Paula and Creed 2004). Mobile plat-
forms could also have contributed to dispersal to the
Gulf of Mexico oil and gas platforms, but that appears
Table 1 Florida Artificial Reefs
with Tubastrea coccinea Name Type Depth Location Date deployed
Donal G McAllister Tug 21 m 26°00.548’ N, 80°05.650’ W 6/23/98
Princess Anne Ship 27 m 26°47.608’ N, 80°00.260’ W 5/23/93
Tenneco Towers Oil rig jackets 32 m 25°58.952’ N, 80°05.100’ W 10/3/85
Tortuga Ship 33 m 25°49.252’ N, 80°05.087’ W 4/95
Rodeo 25 Ship 37 m 26°13.878’ N, 80°03.813’ W 5/12/90
Jay Scutti Tug 20 m 26°09.520’ N, 80°04.760’ W 9/19/86
RSB-1 Ship 35 m 26°13.642’ N, 80°03.896’ W 4/94
Capt. Dan Ship 33 m 26°13.857’ N, 80°03.960’ W 2/20/90
Ancient Mariner Ship 21 m 26°18.117’ N, 80°03.745’ W 6/91
Duane Ship 36 m 24°59.388’ N, 80°22.888’ W 11/28/87
Amesbury Ship 8 m 24°37.391’ N, 81°58.912’ W About 1962
Palm Beach Inlet Floating dock 1 m 26°47.610’ N, 80°02.690’ W NA
Port of Miami
Mitigation Reef
Limestone boulders 11 m 25°44.894’ N, 80°05.683’ W 5/96
Fig. 1 Possible routes of the spread of Tubastraea coccinea in the
Caribbean, Gulf of Mexico, and Florida. Year of the earliest report
in different areas is shown, along with possible routes of spread,
based on dates and current paths. Small arrows present the
generalized current paths in the Caribbean, Gulf of Mexico,
and Florida. Current paths taken from the Ocean Currents web
site (http://oceancurrents.rsmas.miami.edu/caribbean/caribbean-
cs.html), Roberts 1997, and Gittings et al. 1992
506
less likely in Florida. A second species in the genus,
Tubastraea taguensis , has recently been reported to have
invaded Brazil as well, and is easily distinguished in the
field by its yellow color (Figueira de Paula and Creed
2004). In Brazil, both species of Tubastraea have in-
vaded reefs, while in Florida only T. coccinea has been
found. One colony has now been found on the East
Flower Garden Bank, so it is now invading the Flower
Garden Banks National Marine Sanctuary, most likely
from nearby oil and gas platforms. Since it has been in
the western and southern Gulf longer than in the
northern Gulf, it is likely that it is already on reefs in the
southern and western Gulf. It has now been found on a
limestone boulder artificial reef in southeast Florida, so
it is likely that it will soon be found on reefs in Florida,
and reach Bermuda. Artificial structures are clearly
preferred habitat, since in each area they are found first
on artificial structures, and are prolific on some artificial
structures in the Caribbean, Gulf, and Florida. There
are no reports yet of the ecological effects of these
invaders.
The pattern of the spread of T. coccinea in the Wes-
tern Atlantic is very similar to the pattern of spread of
the die-off of the urchin Diadema antillarum in 1983–
1984. The urchin die-off began in Caribbean Panama,
and spread rapidly with the currents to the north and
into the Gulf of Mexico and to Florida. The die-off also
moved eastward from Panama and Florida to the east-
ern Caribbean. Lessios et al. (1984) reported that it was
carried eastward by an eastward-flowing current along
the northern shore of South America. Roberts (1997)
reports that there are weak nearshore counter currents
along most coastlines in the Caribbean. The die-off
spread throughout the Caribbean, Gulf of Mexico, and
to Florida and Bermuda all within 1 year (Lessios et al.
1984; Lessios 1988). In contrast, T. coccinea has required
about 60 years to spread throughout the Caribbean and
Gulf and reach Florida. Presumably this is because the
time between infection of the urchins by the microbe and
the subsequent release of large numbers of new microbes
was probably only a few days (Bauer and Agerfer 1987),
while the time from the settling of T. coccinea larvae
until colonies can release larvae is probably a few years.
Observations in aquaria suggest that larval release
begins at about 18 months age (D. Stettler, personal
communication). If T. coccinea requires about 18
months to begin releasing larvae, and it took 60 times as
long for it to spread across the Western Atlantic as the
urchin die-off, dividing 18 months by 60 might give an
estimate of the time required for the microbe to be re-
leased from infected urchins. The resulting 9 days ap-
pears to at least be the right order of magnitude, which is
consistent with the view that the time to release propa-
gules is a controlling factor in the rate of spread.
Transportation on boat hulls has the potential to have
spread the coral through this area in much less than
60 years, and would likely produce a pattern that would
follow shipping routes instead of current paths. The first
known locations were Puerto Rico and Curacao. Dis-
persal by current from one of these to the other appears
unlikely. However, Puerto Rico was a major coaling
station and shipping port. T. coccinea might have been
brought there first by a ship and carried on to Curac¸ ao
before subsequent dispersal by currents.
Acknowledgements We would like to thank Pamela Fletcher, Keith
Mille, Tim McIntosh, Janet Phipps, Rhett Butler, Julian Sprung,
and Tim Mcintosh for observations of T. coccinea in Florida, and
Emma Hickerson for observations of T. coccinea discovered by
Heidi Lydersen-Bulman on the East Flower Garden Bank. We
thank Rhett Butler for collecting the coral sample, and Daniela
Stettler for aquarium observations.
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