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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
155
STATE OF DEEP CORAL ECOSYSTEMS
IN THE U.S. PACIFIC ISLANDS REGION:
HAWAII AND THE U.S. PACIFIC TERRITORIES
Frank A. Parrish
1
and Amy R. Baco
2
1
Pacic Islands Fisheries Science Center,
NOAA
2570 Dole St. Honolulu, HI 96822
2
Woods Hole Oceanographic Institution
Biology Department
MS#33, 250 Redeld
Woods Hole, MA 02543
I. INTRODUCTION
The U.S. Pacic Islands Region consists of
more than 50 oceanic islands, including two
archipelagos (Hawaii and Mariana Islands), parts
of four other archipelagos (Samoa, Line Islands,
Phoenix Islands, and Marshall Islands), and
numerous seamounts in proximity to each of these
groups. These islands include the State of Hawaii,
the Commonwealth of the Northern Mariana
Islands (CNMI), and the territories of Guam and
American Samoa, as well as nine sovereign
Federal territories—Midway Atoll, Johnston Atoll,
Kingman Reef, Palmyra Atoll, Jarvis Island,
Howland Island, Baker Island, Rose Atoll, and
Wake Island). This area also encompasses the
Pacic Island States in free association with the
United States (former U.S. trust territories also
known as the Freely Associated States) including
the Republic of Palau, the Federated States
of Micronesia (Chuuk, Pohnpei, Kosrae, and
Yap), and the Republic of the Marshall Islands.
This region includes some of the most remote,
unpopulated islands in the Pacic, as well as
many densely populated islands, and it extends
from the South Pacic (e.g., American Samoa;
14º S latitude) to the North Pacic (Kure Atoll 28º
N latitude) (Figure 4.1). The punctuated habitat of
the Pacic Region distinguishes deepwater coral
communities biogeographically and ecologically
from other areas in the United States. Because
of the isolated nature of these islands (especially
Hawaii and the Northwestern Hawaiian Islands),
they possess some of the highest levels of marine
endemism recorded anywhere on earth.
While trace coral samples from anecdotal
dredging and bycatch suggest a wide distribution
of deep corals throughout the Pacic, the only
detailed assessment of deep corals within the U.S.
waters of the Pacic has been in the Hawaiian
Archipelago. Antipatharians were rst reported
from Hawaiian waters more than 75 years ago
(Verrill 1928). The earliest descriptions of deep
octocorals in Hawaii are recorded by Dana (1846),
with Nutting (1908) reporting 68 species. Other
signicant contributions to the species lists of this
region include Muzik (1979) and Grigg and Bayer
(1976) for octocorals, as well as Vaughan (1907)
and Cairns (1984, 2006) for scleractinians. Wells
(1954) provides data on the Marshall Islands.
Pacic deep coral research has expanded greatly
over the last four decades, primarily as a result
of the establishment of commercial sheries for
black, pink, and gold coral off the main Hawaiian
Islands, and subsequent development of shery
management plans for these resources by the
State of Hawaii and the Western Pacic Fishery
Management Council. Deep corals are harvested
as raw material for the jewelry trade. The coral
supports a portion of a $70 million Hawaii-based
industry that employs roughly 650 people in its
manufacturing facility and 50 retail stores (Carl
Marsh—Maui Divers pers. comm.)
Commercial beds of black coral were rst
discovered at a depth of 30-75 m off Lahaina,
Maui in 1958. Some of the earliest ecological
work on black corals was carried out in the 1960s
in the channel waters off Maui using SCUBA
(Grigg 1965). The Maui black coral bed has
remained the focus of coral harvesters throughout
the shery’s history and has been periodically
resurveyed over the last 40 years. These studies
have provided the longest data sets available
worldwide on the status and trends of black coral
populations and the effects of the commercial
shery and other natural and anthropogenic
stressors. In the mid 1960s, isolated patches
of pink (Corallium spp.), gold (Gerardia sp.)
and bamboo (Lepidisis sp., Acanella sp.) corals
PACIFIC ISLANDS
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
were identied at 300–500 m depths north of
Midway Island (Milwaukee Banks) and off Oahu
(Makapuu Bed) (Grigg 1993). A long-term deep
coral research program focused on precious
corals began at the University of Hawaii in 1970.
Many of the earliest surveys of precious coral beds
used tangle net dredges and other nonselective
gear. A key advancement in Hawaii’s deep coral
research infrastructure was access to the two-
person submersible Deep Star 2 from General
Dynamics. Aside from periodic research (Grigg
1993) the sub was leased to commercially harvest
coral for the shery between 1974 and 1979. In
1980 the submersible was renamed the Makalii
and became the centerpiece of the newly formed
Hawaii Undersea Research Laboratory (HURL),
an established node of NOAA’s Undersea
Research Program (NURP). The facility has
since expanded, replacing the Makalii with two,
deep-diving 3-person submersibles (Pisces IV
and Pisces V) and a dedicated support vessel
equipped with a multibeam bottom mapper and
a remotely operated vehicle (RCV-150) (Chave
and Malahoff 1998). This new infrastructure
expanded the focus of coral research and
increased participation by more researchers.
This chapter provides a summary of what is
known about deep corals within the Pacic
Islands Region. In keeping with the intent of this
national report, the chapter will mostly focus on
corals deeper than 50 m. However, shallower
black corals will be included. Most of the
information available on black corals, precious
corals, and other deep corals are from the
Hawaiian Archipelago, where most of the surveys
have been conducted. Studies have focused on
the taxonomic and genetic composition of the
region’s coral community, ecological relationships
between corals and other organisms, and on the
distribution and dynamics of deep corals. Much
of this work is focused on the coral taxa that are
targets for the commercial shery. Also discussed
are the measures that have been employed to
Figure 4.1. Map of the Pacic Basin showing U.S. islands and their Exclusive Economic Zone that comprise
the Pacic Islands Region for the National Marine Fisheries Service.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
157
protect deep coral ecosystems and to manage
the commercial shery.
II. GEOLOGICAL SETTING
The U.S. Pacic Islands lack the shelf area that
typically denes the deep-sea benthic habitats
of the continental United States. Instead, the
submerged slopes of volcanic pinnacles that rise
steeply from abyssal depths of 4–7 km provide
the hard substratum that deep corals colonize.
The region has endured a long history of plate
drift, subsidence, and sea level rise, and many
of the volcanic islands have drowned creating
numerous submerged banks and seamounts. A
striking feature of the Pacic Basin is the linear
nature of the island chains and seamounts. They
are aligned in a north-northwesterly direction,
a consequence of the northwesterly drift of the
Pacic plate over stationary volcanic hotspots
(Kennett 1982). The resulting islands and
seamounts are progressively older in proportion
to distance from a hotspot. For example, the
island of Hawaii lies above the mantle plume and
is the only island in the Hawaiian Archipelago that
is volcanically active. To the northwest, volcanism
on Oahu ceased about three million years ago;
Kauai about ve million years ago; and Midway
Island about 27 million years ago (Grigg 1988a).
This geologic process denes the Pacic plate
and, as a result, the Pacic Islands region has
some of the youngest (main Hawaiian Islands)
and oldest (Line Islands) volcanic archipelagos
in the world.
III. OCEANOGRAPHIC SETTING
The Pacic is composed of two large gyres,
the northern and southern central gyres. In the
South Pacic, southeast trade winds drive the
South Equatorial Current westerly between 15
o
S and 3
o
N latitude. Within the South Equatorial
Current is the Cromwell Current, or Pacic
Equatorial Undercurrent. This current exists at
depths of 70–200 m, and is approximately 450
km wide and ows with velocities of up to 5 km h
-1
for a distance of up to 13,000 km in the opposite
direction of the South Equatorial Current (Tchernia
1980; Thurman 1981). In the North Pacic, the
North Equatorial Current ows westward at 1 km
h
-1
between 8
o
and 20
o
N latitude. The Equatorial
Counter Current is located between the North
Equatorial Current and the South Equatorial
Current and travels eastward at slightly more
than 2 km h
-1
(Thurman 1981). The boundaries
of these water masses overlap and contribute
to long-distance dispersal of pelagic larvae,
particularly in the western Pacic. The northern
Hawaiian seamounts fall in the northern portions
of the north gyre; Hawaii, Wake and Johnston are
in the center of the North Pacic gyre; Kingman
and Palmyra are in the equatorial/eastern Pacic;
Jarvis, Howland, and Baker are in the equatorial
system; American Samoa is in the equatorial
portion of the southern gyre; and the Mariana
Islands are affected by the north central gyre,
the equatorial systems, and the Kuroshio current
from Asia.
While deep water masses originate from surface
currents, no deep water masses form in the
Pacic Basin. Deep water migrates in from
the Atlantic via the southern hemisphere with a
uniform temperature and salinity below about
2000 m (Knauss 1996). The deep water ows
northward at depths below 2500 m and southward
above 2500 m. Seamounts, pinnacles, and other
structures obstruct current ow and can generate
eddies of varying intensity, depending on the
current velocity, depth or height of the seamount,
slope of the side, and strength of the seawater
stratication. Both cold and warm water eddies
are formed as a result of a seamount obstructing
current ow in the deep ocean. Typically,
anticyclonic (cold water) eddies are formed above
the seamount and remain tightly associated
with the top of the seamount, while the cyclonic
(warm water) eddy is formed downstream
behind the seamount (Kamenkovich et al.
1986). Deep corals are thought to benet from
the ow acceleration, larval retention, and high
nutrient waters from deep upwelling caused by
the presence of the seamount and the generated
eddies (Genin et al. 1986; Mullineaux and Mills
1997; Coutis and Middleton 2002).
Oxygen in the deep water of the Pacic has been
depleted to very low levels as a result of the length
of time it takes for water to move into and across
the Pacic Basin. Having aged thousands of
years, oxygen averages 0.5–4.5 ml l
-1
versus the
signicantly higher Atlantic average of 3.0–6.5
ml l
-1
(Thurman 1981). The effect of low oxygen
levels on deep corals is poorly documented;
however, Wishner et al. (1990) attributed patterns
in the abundance and distribution of sponges,
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
sea pens, and other benthic organisms to depth-
specic patterns in oxygen levels.
IV. STRUCTURE AND HABITAT-FORMING
DEEP CORALS
Most of the major deep coral groups are known
to exist in the U.S. Pacic region. However,
most species have been identied only around
the Hawaiian Archipelago largely because deep
waters around other U.S Pacic Islands have not
yet been explored. Published records of deep
corals from the Hawaiian Archipelago include more
than 137 species of gorgonian octocorals and 63
species of azooxanthellate scleractinians, with
21% of the scleractinians thought to be endemic
to Hawaii (Cairns 2006, See chapter appendix).
A 2003 cruise in the Northwestern Hawaiian
Islands (A. Baco, unpublished data) identied
eight new species of octocorals, two new genera
and several new species of antipatharians, three
new stylasterid species (Cairns 2005), and 1
new zoanthid species. In addition, a 2004 main
Hawaiian Islands cruise collected at least three
new species of octocorals, two new species of
Taxa
Reef-
Building Abundance
Maximum
Colony
Size Morphology
Associations
with Other
Structure-
Forming
Invertebrates
Colony
Spatial
Dispersion
Overall
Rating of
Structural
Importance
Enallopsammia
rostrata No Medium Medium Branching Many Clumped Medium
Other
scleractinians No Low Small
Non-
Branching Many Solitary Low
Gerardia sp. No High Large Branching Many Clumped High
Shallower
antipatharians No Low Large Branching Few Clumped High
Other
octocorals and
antipatharians in
deeper water No High Med Branching Many Clumped High
Other
octocorals and
antipatharians
in precious coral
beds No Medium
Medium
-Low Branching Many Clumped High
Corallium
secundum No High Medium Branching Many Clumped Medium
Corallium
laauense No High Medium Branching Many Clumped Medium
Isidids in deeper
water No High Med Branching Many Clumped Medium
Table 4.1 Structure-forming attributes of deep corals in Hawaii.
Table Key
Attribute
Measure
Reef-Building
Yes/No
Relative Abundance
Low/ Medium/ High
Size (width or height)
Small (<30cm)/ Medium (30cm-1m)/ Large (>1m)
Morphology
Branching/ Non-branching
Associations
None/ Few (1-2)/ Many (>2)
Spatial Dispersion
Solitary/ Clumped
Overall Rating
Low/ Medium/ High
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
159
antipatharians, and provided range extensions
for several genera and species of corals that
were not previously known from Hawaii. Thus,
although an extensive species list exists for the
Hawaiian Archipelago, the high rate of discovery
of new species and new records implies the
Archipelago is also largely undersampled.
In general, the deep corals in the Pacic Islands
do not form the extensive reef structures observed
in the Atlantic and South Pacic. Instead, corals
grow attached directly to the exposed fossil
carbonate, basalt or manganese substratum.
Octocorals and antipatharians have been found
to grow in high densities at numerous sites,
particularly on summit areas of seamounts or
other topographic highs, where they often form
extensive coral gardens or “beds” with abundant
associated invertebrates. The coral taxa listed in
this chapter are those that present a conspicuous
relief prole that could serve as a source of
habitat (Table 4.1). At death, these taxa decay
from physical and bioerosion forces until they
detach from the substratum and are swept away.
Cemented reefs from accumulated dead material
have not been observed.
a. Stony corals (Class Anthozoa, Order
Scleractinia)
Enallopsammia rostrata is an arborescent
scleractinian coral in the Family Dendrophyllidae.
The full depth range for this species is listed as
229–2165 m in Cairns (1984), but it has been
observed in Hawaii primarily at depths of 500–
600 m. In some areas it forms bushy colonies,
with dead coral accumulating near the base
of the colony much like that observed among
Lophelia reefs in the Atlantic. Further exploration
and characterization of this species is needed to
determine its abundance throughout the region
and its potential role in forming habitat.
Madrepora kauaiensis and M. oculata also
occur in Hawaii and have the potential to form
reef structures, however, little is known of their
abundance or distribution in the Archipelago.
Besides these examples, scleractinians that
have been observed are primarily solitary cup
corals. They can occur in abundance, e.g., on
Cross Seamount (A. Baco pers. obs), but many
species are small and not observable with a
submersible, preventing a true determination of
their distribution. A complete species list (to date)
for Hawaii can be found in Cairns (2006) and is
also included in the Appendix to this chapter.
b. Black corals (Class Anthozoa, Order
Antipatharia)
Fourteen genera of black corals are reported from
the Hawaii-Pacic region with species found in
both shallow and deep habitats. The shallowest
genera (Cirripathes spp. and Antipathes spp.)
prefer shaded or low light areas and can occur
underneath ledges and in caves in shallow water
(e.g., Cirrhipathes anguina can occur in 4 m depth)
where surge is minimal, or in the open on steep
walls at deeper depths. Antipathes spp. appears
to settle predominantly in depressions, cracks or
other rugged features along steep ledges, with
few colonies found on smooth basaltic substratum
(Grigg 1965). Shallower antipatharians in
Hawaii also appear to prefer substrates that are
encrusted with calcium carbonate from coralline
algae, bryozoans, and corals. The highest
densities are found on hard sloping substratum, in
areas with 0.5–2 knot currents (Grigg 1965). The
best studied black corals are the commercially
harvested species Antipathes dichotoma and
Antipathes grandis. Recent taxonomic work
(D. Opresko pers. comm.) on the Hawaiian A.
dichotoma suggests it is a new species and is
currently being referred to as Antipathes cf.
curvata. The A. cf. curvata and A. grandis exhibit
similar growth rates (6.42 cm yr
-1
and 6.12 cm yr
-1
,
respectively) and reach reproductive maturity at
ages 12–13. Fertilization takes place externally
in the water column, and light and temperature
inuence larval settlement patterns. The larvae
of A. cf. curvata and A. grandis are negatively
phototactic, and the lower depth limit coincides
with the top of the thermocline (~ 100 m) in the
main Hawaiian Islands (Grigg 1993).
Much less is known about deeper genera of
black corals. They have similar morphologies to
the shallower corals, including colonies shaped
as whip-like laments (Stichopathes spp.) and
as branching, sometimes “feathery” colonies
(Myriopathes, Bathypathes spp., Stauropathes
and Leiopathes). However, the life history of
these deeper genera is likely to be much different
than their shallower relatives. Radiometric dating
of A. cf. cruvata and Leiopathes indicates the
deeper Leiopathes genera grow 10 to 70 times
slower than the shallower A. cf. curvata (Roark
et al. 2006).
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
c. Gold coral (Class Anthozoa, Order
Zoanthidea)
No species of zoanthid has yet been described
from deep water in Hawaii although taxonomically,
at least six species have been observed
and,ollected (Baco, unpublished data). The gold
coral, Gerardia sp., is probably the most common
and certainly the largest of these species. It has
an arborescent morphology similar to gorgonians,
and colonies have been observed as tall as 2–3
m in height (Figure 4.2 B). Gerardia sp. is widely
distributed throughout the Hawaiian Archipelago
and into the Emperor Seamount Chain at depths
of 350–600 m.
Zoanthids in Hawaii have been observed to
colonize other living coral species as well as
hard bottoms. In the case of Gerardia sp., the
zoanthids may eventually replace the host colony
completely. It is not known if Gerardia sp. can
outcompete the living tissue of the host or if it
opportunistically colonizes and spreads after a
portion of the host coral has been decorticated
by predatory urchins or some other cause. The
life span of Gerardia sp. is uncertain. Counts of
growth bands assumed to be annual in periodicity
have provided an estimated lifespan of around 40
years (Grigg 2002). Recent radiometric work on
the Hawaiian species has estimated the life span
of gold coral samples between 450 and 2700
years (Roark et al. 2006), which is consistent
with ndings from radiometric aging on Gerardia
sp. in the Atlantic (Druffel et al. 1995).
Gold coral was one of the few corals seen during
the 2005 Line Island surveys and it was present
at Jarvis, Palymra, and Kingman. However, the
colonies were sparse, with no patches large
enough to be called a “bed.” All colonies were
infested with other unidentied zoanthids. Surveys
at the base of the cliffs below the spot where the
individual gold colonies were attached found no
accumulation of fallen colonies, suggesting gold
coral has always been in low abundance in the
region (Frank Parrish pers. obs.).
d. Gorgonians (Class Anthozoa, Order
Gorgonacea)
Gorgonian octocorals are by far the most
abundant and diverse corals in the Hawaiian
Archipelago. Two species, Corallium laauense
(red coral; formely identied as Corallium regale)
and Corallium secundum (pink coral) are known
to occur at depths of 350–600 m on islands and
seamounts throughout the Hawaiian Archipelago
(Grigg 1974, 1993; Baco, unpublished data)
and into the Emperor Seamount Chain (Bayer
1956). Growing to more that 30 cm in height
the Corallium spp. occur in a variety of red/pink
color shades, and the height and shape of the
colony’s “fan” can vary considerably (Figure
4.2 A). They are often found in large beds and
usually support a high diversity of invertebrates
with an abundance of other octocorals, zoanthids,
and sometimes scleractinians co-occuring in
the beds. C. secundum and C. laauense are
gonochoristic (Grigg 1993; Waller and Baco in
press) and are estimated to reach reproductive
Figure 4.2. Photos of the two primary Genera that comprise the Hawaiian precious coral shery
A—Corallium sp. and B—Gerardia sp. Photo credit F. Parrish, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
161
maturity at 12–13 years (Grigg 1993). These
species are relatively long lived, with some of the
oldest colonies observed within Makapuu Bed
about 0.7 m in height and approximately 80 years
old (Grigg 1988b, Roark 2006). Populations of
C. secundum appear to be recruitment limited,
although in favorable environments (e.g.,
Makapuu Bed) populations are relatively stable,
suggesting that recruitment and mortality are in a
steady state (Grigg 1993).
More than 130 other species of octocorals are
known from the Archipelago and they represent
a diverse array of families and genera. Most
abundant are the Families Coralliidae, Isididae,
Primnoidae, Plexauridae, Chrysogorgiidae and
Acanthogorgiidae. To our knowledge, besides
the species discussed above, there isn’t any
information on the biology and ecology of these
groups in this region. Further discussion of their
depth distributions are in the following sections
and a complete species list (to date) is included
in the Appendix to this chapter.
e. True soft corals (Class Anthozoa, Order
Alcyonacea)
The Alcyonacea are represented in this region by
only 12 species in three families. Of these, the
genus Anthomastus is the most widely distributed.
It is often observed in precious coral beds in
patches with large number of small individuals
surrounding a larger individual (A. Baco pers.
obs.). The bright purple Clavularia grandiora
has been observed growing on Gerardia at a
number of sites throughout the Archipelago (A.
Baco pers. obs.)
Like the gorgonians, little else is known about
the biology and ecology of these species in
this region. A complete species list (to date) is
included in the Appendix to this chapter.
f. Pennatulaceans (Class Anthozoa, Order
Pennatulacea)
Pennatulaceans seen in Hawaii tend to be more
abundant in areas high in sediment, although
Kophobelemnon sp. has occasionally been
observed on adjacent hard bottoms near the
Cross Seamount deep coral bed. Near the
Keahole deep coral bed on the island of Hawaii,
a single species (as yet unidentied) occurs in
high abundance in patches of sediment at depths
of about 400 m (A. Baco pers. obs.). Again, little
else is known about the biology and ecology of
these species in this region. A complete species
list (to date) is included in the Appendix to this
chapter.
g. Stylasterids (Class Hydrozoa,
Order Anthoathecatae)
Four species of stylaserids are present in
Hawaii, but they are distributed very patchily
throughout the Archipelago. An extreme example
is Disticophora anceps. It has a very wide depth
range but has only been found on the northwest
slope of Laysan Island in densities of several
colonies per square meter in some areas (Cairns
2005; A. Baco unpublished data). Again, little
else is known about the biology and ecology of
this group in this region.
Hawaiian stylasterids are discussed in Cairns
(2005) and a complete species list (to date) is
included in the Appendix to this chapter.
V. SPATIAL DISTRIBUTION OF CORAL
SPECIES AND HABITAT
General distribution
Our knowledge of the spatial distribution of deep
corals in the U.S. Insular Pacic is limited to
Hawaii. Even in Hawaii, very little of the deep
sea has been explored and every research
expedition is yielding large numbers of new
species. Until 2003, the majority of studies in
Hawaii came from sparse trawl data or had
concentrated on the harvested black, gold, and
pink corals. Often referred to as “precious corals,”
these are the primary deep coral taxa harvested
for the jewelry trade. Most of these are found
between depths of 300 and 500 m and have
been collected by dredge or submersible. Often,
black coral (Antipathes spp.) is distinguished
from the rest of the precious corals because the
Antipathes taxa that are used for jewelry occur
much shallower (<100 m) and are harvested
by scuba divers. Beside precious corals, many
other taxa of deep corals have not been studied
because they were not one of the management
unit species of the precious coral shery. In 2003
and 2004, the Pisces submersibles were used
for the rst studies of non-precious corals, thus
extending the exploration of corals well below
previously surveyed depths. In 2005, the rst
surveys conducted outside of Hawaii using the
Pisces submersibles were at Rose Atoll and the
U.S Line Islands (Jarvis, Palmyra, and Kingman).
PACIFIC ISLANDS
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
The distribution of deep corals in the rest of the
U.S. Pacic is unknown.
Depth clearly inuences the distribution of
different coral taxa and certainly there is
patchiness associated with the presence of
premium substrate and environmental conditions
(ow, particulate load, etc.). The environmental
suitability for colonization and growth is likely to
differ among coral taxa. For example, Corallium
secundum appears to grow in large numbers
in areas of high ow over carbonate pavement;
Corallium laauense grows in an intermediate
relief of outcrops; and Gerardia sp. grows in high
relief areas on pinnacles, walls, and cliffs (Parrish
in press). These habitat differences may reect
preferred ow regimes for the different corals
(e.g., laminar ow for C. secundum, alternating
ow for Gerardia sp.).
Black coral beds
Black coral beds are found off the main Hawaiian
Islands at depths of about 30–110 m. The largest
bed covers an estimated area of 1.7 km
2
and lies
in the middle of the Auau Channel, between Maui
and Lanai, encrusting a drowned land bridge
between the two islands at depths of 30-90 m
(Grigg et al. 2002). A smaller black coral bed
(0.4 km
2
) is located off Kauai and another at the
southern end of the island of Hawaii (Figure 4.3).
The dominant species found in these locations
are Antipathes cf. curvata (95% of the population)
followed by Antipathes grandis. Grigg (1976)
estimated a total standing crop of A. cf. curvata
for the Auau Channel area, between 40 and 70
m, to be 166,000 kg or 84,000 colonies, while the
bed at Kauai contained 40,000 kg. Myriopathes
ulex is found in deeper locations (110–565 m)
off the main Hawaiian Islands, along with other
species of antipatharians absent from shallower
depths (Devaney and Eldridge 1977) (see
chapter appendix). Little commercially important
black coral has been found in the Northwestern
Hawaiian Islands (Grigg 1974), perhaps due to the
shoaling of the thermocline towards the northwest
end of the chain. Other species of black corals
Figure 4.3. Topographic map of the main Hawaiian Islands with the three known black coral beds marked.
Inset shows a diver conducting a coral survey. Map creidt: F. Parrish, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
163
occur in the Northwestern Hawaiian Islands and
include taxa such as Cirripathes, Stichopathes,
Stauropathes, Bathypathes, Myriopathes ulex,
Trissopathes, Umbellopathes, Dendropathes,
and Leiopathes.
Between Black and Precious Coral Beds
The depth zone between the black coral beds
and the precious coral beds has had less study.
Corals have been observed in this zone; for
example, an abundance of octocorals occur at
the Makapuu coral bed on the island of Oahu,
shallower than the precious corals, but they have
not been sufciently sampled to comment on
diversity or species composition.
Precious coral beds
Probably the most abundant of Hawaii’s known
deep corals are the precious corals, including
octocorals Corallium laauense (red coral) and
Corallium secundum (pink coral), and the
zoanthid Gerardia sp. (gold coral). These species
are known to occur in signicant abundance in
at least 16 locations in the Hawaiian Archipelago
at depths of 350–600 m (Grigg 1974, 1993;
Baco, unpublished data) and into the Emperor
Seamount Chain (Bayer 1956). Within a given
coral bed, the two primary genera (Corallium
and Gerardia) are usually found, but the ratio
of abundance can vary greatly (Parrish in
press)(Figure 4.4). It is difcult to estimate the
size of coral beds, so only relative differences in
bed size (based on impressions of coral density
and the area covered by the submersible track)
were shown in Figure 4.4 to determine the size of
pie diagrams. Most precious coral sites also have
a number of other noncommercial taxa; these
include various octocorals (e.g., Callogorgia,
Paracalyptrophora, Acanthogorgia, Lepidisis,
Keratoisis, Isidella, Kereoides, Paragorgia, and
various paramuriceids) and antipatharians (e.g.
Leiopathes, Trissopathes, and Bathypathes) (A.
Baco, unpublished data).
Of the known coral beds, the Makapuu coral bed
is the best studied and most diverse. It is located
between 375 and 450 m depth in the channel
between the islands of Oahu and Molokai. The
bed comprises an area of about 3.6 km
2
, with
the most abundant coral C. secundum, at a
mean density of 0.22 colonies per square meter
between 365 and 400 m (Grigg 1988b). Other
corals found at Makapuu include bamboo coral
(Lepidis olapa, 0.041 colonies m
-2
; Acanella spp.,
0.001 colonies m
-2
), gold coral (Gerardia sp.,
0.0005 m
-2
), as well as three genera of gorgonians
Figure 4.4. Topographic map of the Northwestern Hawaiian Islands with coral survey dive sites of the sub-
mersible Pisces V (inset). Pie charts represent the relative amount of coral among sites and the within-site
fraction of the two primary precious coral genera. 3D map and Pisces photo credit: Hawaii Undersea Research
Laboratory.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
(Narella sp., Psuedothesea sp and Callorgorgia
gilberti), a sea pen (Stylatula sp.), and black coral
(Leiopathes n. sp.). Many taxa of no commercial
interest are also present in or adjacent to the bed
including Enallopsammia rostrata, Thouarella
hilgendor, acanthogorgiids, Paragorgia sp.,
Paramuriceidae, Trissopathes pseudotristicha
(antipatharian) and a number of undescribed
octocorals.
Beyond Precious Coral Beds
Explorations have been conducted at only a
few sites below precious coral depths; Pioneer
Ridge, the small seamount southeast of Laysan
Island, an unnamed seamount east of Necker
Island, Cross Seamount, and Keahole Point. At
these deeper depths in high current areas such
as ridges and pinnacles, a fair amount of overlap
appears to occur in species composition of both
corals and sponges between sites. Although
the number of observations is very limited, there
appears to be a transition in species below about
600 m, from Corallium- and Gerardia-dominated
communities, to a different suite of species. Many
species of chrysogorgiids, primnoids, isidids,
coralliids, and antipatharians begin to appear
around this depth (A. Baco, unpublished data).
Among the more common octocoral genera
observed are: Chrysogorgia, Metallogorgia,
Iridigorgia, Narella, Calyptrophora, Candidella,
Keratoisis, Isidella, Acanella, Corallium, and
Paragorgia, as well as the antipatharian genus
Bathypathes. The depth distribution of many,
but not all, of these species appears to continue
below 1800 m (Baco, unpublished data).
V. SPECIES ASSOCIATIONS WITH DEEP
CORAL COMMUNITIES
The harvesting of deep corals has prompted
a number of studies focused on the species
associations with deep coral communities.
Nationally, most effects to coral-associated
species come from trawling and dredging,
which are banned in U.S. Pacic waters. The
harvesting of precious corals is allowed using
selective methodologies such as hand collection
by scuba divers or using the manipulator of a
submersible. Take of deep corals as a shery
target is a direct effect to the bottom habitat with
uncertain ecological consequences. Studies have
been conducted to address the NOAA mandate
of essential sh habitat, protected species, and
ecosystem concerns. These have historically
been focused on sh and only recently have been
expanded to include invertebrates.
Commercial shery species
With the exception of the shery that harvests
precious corals, there is little evidence of a direct
association between precious corals and other
shery targets. However, these evaluations
have been limited to comparing the overlap in
depth ranges and making infrequent underwater
observations. Even less is known about the deep
sea corals not targeted in
the precious coral shery,
or any indirect ecological
effects that any of these
corals may contribute to
commercial shery stocks.
Some of the shallow
coral reef sh targeted by
recreational shers and the
aquarium trade range into
depths where black corals
(antipatharians) can be
found (30–100 m) (Moftt
et al. 1989; Parrish and
Boland 2004; Boland and
Parrish 2005). One of the
commercially sought bottom
sh Aprion virescens (grey
snapper), also lives in this
depth range but most of
the commercial bottom
Figure 4.5. Two species of black coral trees Antipathes grandis (left) and
Antipathes. cf. curvata (right). Photo credit F. Parrish, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
165
sh reside at depths below antipatharians and
above the precious corals (<300 m) (Uchida
and Tagami 1984). The shallowest members of
this group, such as Pristipomoides lamentosus
(pink snapper), Pristipomoides zonatus (oblique-
banded snapper), and Epinephelus quernus
(Hawaiian grouper), have been seen in the
vicinity of deeper black coral trees (Moftt et al.
1989). Similarly, the deeper members including
Etelis carbunculus (ruby snapper) and Etelis
coruscans (ame snapper) have been seen
among the shallower precious corals (Kelley et
al. 2006). The groundsh, Pseudopentaceros
wheeleri (armorhead) and Beryx sp. (alfonsino)
occur throughout this depth range (250–350)
but are more common on the seamounts at the
northern end of the Hawaiian Archipelago (Uchida
and Tagami 1984). There is no information on
the degree of overlap of these sh with deep sea
corals. Heterocarpus sp. (deep-water shrimp)
has been seen among the precious corals but at
densities consistent with other bottom relief types.
Heterocarpus sp. is the focus of an intermittent
main Hawaiian Island trap shery that targets
depths of 500 to 900 m (Moftt and Parrish 1992)
overlapping the lower limit of precious coral
depths, but in the depth range of many other
deep coral species.
Noncommercial species
Fish
Studies of sh associations with deep corals have
focused almost exclusively on the inventory of sh
taxa and appraisal of the obligate or facultative
roles corals play in sh assemblages. The sh
community of the Auau Channel black coral bed
was recently surveyed (Boland and Parrish 2005)
and 95% of the sh found in and around the
black corals were known to commonly occur on
shallower reefs where black corals do not grow.
Oxycirrhites typus (the longnose hawksh), was
found exclusively within the black coral trees.
Behavioral data indicated that although most of
the reef sh routinely passed through the coral
Figure 4.6. Examples of invertebrates found associated with deep corals (A-basket star, B-ophiuroid,
C-asteroid. Photo credit A.Baco, WHOI.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
branches, only four species reliably used the
corals for cover when evading a threat (Figure
4.5). Although there is little or no known obligate
relationship between sh and black coral, the
coral colonies contribute to the sh community
by enhancing the vertical aspects of the deep
reef and perhaps improving the corridors for sh
movement.
Most of the sh of the deep slope and subphotic
depths are noncommercial species (Chave and
Mundy 1994). The surveys of sh communities
at deeper subphotic depths indicated few sh
associations with precious corals (Parrish 2006).
Many of the 42 sh taxa observed were seen
to use Gerardia and Corallium spp. as shelter
interchangeably with abiotic relief sources.
Species richness of sh was not observed to differ
between areas with corals and those without.
Most sh taxa were observed in low numbers
with only a couple of dominant species. Areas
with corals often supported greater sh density,
but statistical evaluations suggested that this
was based on co-occurrence of sh and coral in
areas of relief and high ow and not based on a
dependency of sh on corals. Also, differences
were not seen in the mean size of sh in or outside
of the coral beds. Of the three commercial coral
species, sh oriented mostly around Gerardia sp.
probably because it is signicantly taller than the
two Corallium species. Classifying the sh into
functional groups revealed “benthic hoverers”
as the segment of the sh community that most
frequently used Gerardia sp. as shelter.
Although use of corals by the sh appeared
incidental, it is important to consider these studies
were conducted in summer, during the day and
focused exclusively on adult sh. It is unknown
if the corals play some seasonal or diurnal role in
the sh ecology or if juvenile stages rely on the
coral colonies.
Invertebrates
There is a wide array of invertebrate species
associated with deep corals. Black corals are
Figure 4.7. The
purple octocoral
Clavularia gran-
diora growing
on a Gerardia sp.
colony.Photo credit
A. Baco, WHOI.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
167
known to be colonized by oysters, bryozoans,
and shrimp (Hoover 1998). The most common
invertebrates associated with the deeper precious
corals include zoanthids, anemones, galathaeoid
crabs, sponges, ophiuroids, and basket stars
(Figure 4.6). Gerardia sp. is perhaps the best
known coral that overgrows the skeletons of
other coral species. Examination of Gerardia sp.
skeletons and many submersible observations
suggests it has an obligate need to start its
colony over the skeleton of other corals. Bamboo
corals seem to be the most frequent target,
particularly Isidella trichotoma, but Gerardia has
been observed growing on quite a number of
species. Several other species of unidentied
zoanthids also appear to prefer to grow on other
corals, including Gerardia sp. At least one of
these zoanthids was observed growing on basalt
substrate as well as encrusting other corals (A.
Baco pers. obs.). Octocorals also can grow
on the skeletons of other coral species. The
bright purple Clavularia grandiora has been
observed growing on Gerardia at a number of
sites throughout the Archipelago (Figure 4.7).
In all of these cases, it is not clear whether the
overgrowing corals are actually killing or injuring
the coral whose skeleton they are growing on, or
if they have simply colonized a dead area of the
skeleton.
Many coral colonies have a number of commensal
invertebrates such as galathaeoid crabs,
barnacles, sponges, anemones, polychaetes,
crinoids, ophiuroids, and basket stars (Chave and
Malahoff 1998; A. Baco and Shirley, unpublished
data). Casual observations suggest some of these
commensals may be specic to their host while
others are more generalized. The relationships
between the coral and the commensal, e.g.,
symbiotic, parasitic, obligate, facultative, etc.,
have not been determined in most cases and
many of the commensal species have yet to be
identied. One of the more notable commensal
relationships is the general association of
polychaetes with species in the genus Corallium.
Each Corallium species appears to have its own
species of polynoid polychaete. In Corallium
secundum and Corallium laauense, these
polychaetes can reach fairly high densities. The
polychaetes live in tunnels under the coral soft
tissue with the skeleton often growing over the
polychaete tunnels. This relationship has also
been observed in other corals such as Candidella
helminthophora.
A more generalized commensal is the unbranched
basket star, Asteroschema. Asteroschema sp.
has been observed in a number of different species
of octocorals at a number of sites, but not on the
surrounding substratum. Dead coral skeletons
also appear to provide good recruitment habitat
for many invertebrate species. Many types of
sessile fauna have been observed as well as
several types of young corals. In particular, young
colonies of the red coral, Corallium laauense
have been observed growing on dead skeletons
of Gerardia sp.
Besides the galathaeoid crabs that inhabit the
branches of coral colonies, a number of larger
crabs are routinely encountered patrolling the
bottom around deep corals (e.g., crabs in the
families Homolidae, Parapaguridae).
Predation on deep corals by resident invertebrates
also occurs. Seastars feed on coral colonies
by everting their gut, leaving behind patches
of bare coral skeleton. Cidarid urchins are
also known to feed on deep corals and these
urchins have been observed on deep corals
in Hawaii. However, an absence of bare coral
skeleton around the urchin’s location suggests it
is unlikely they are consuming the coral tissues
(A. Baco, unpublished data). A single species of
orange crinoid was observed in 2004 in very high
densities at the Makapuu coral bed (A. Baco,
unpublished data). Grigg 2002 comments on an
abundant crinoid observed during night dives at
this site. The crinoids now cover many octocoral
colonies and anything else that sticks up more
than a few millimeters off the bottom (A. Baco,
unpublished data), suggesting their abundance
has increased over Grigg’s observations. Their
origin and the reason for their recent increase in
population density, as well as their potential for
competing with corals for food, are unknown.
Monk seals
In the lower Northwestern Hawaiian Islands,
the endangered Hawaiian monk seal has been
documented to routinely visit deep corals as
part of its foraging activities. However, there
have been no reported interactions between
monk seals and the precious coral harvesting
in the main Hawaiian Islands. Telemetry and
scatological analysis indicate seals prey on
bottom-dwelling sh (Goodman-Lowe 1998;
Parrish and Abernathy 2006; Longnecker et
al. 2006). Video cameras tted to seals in the
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
Table 4.2 Potential effects of shing gears on deep coral habitat in Hawaii. Ratings detailed in table key. Tuna/
Swordsh includes: Albacore, Bigeye tuna, Yellown Tuna, Skipjack Tuna, Swordsh, Striped Marlin, Pacic
Blue Marlin, Black Marlin Sailsh, Shortbill spearsh, Wahoo, Dolphinsh, Opah, Pomfret, sharks. Deepwater
shrimp include: two species of Heterocarpus, Snappers and Groupers include: Pink snapper, Flower snapper,
Squirrelsh snapper, Hawaiian Grouper, Ruby-colored snapper, Blue-green snappersh.
Gear Type
Current
Fishery
Use in
Region
Potential
Severity
of Impact
Potential
Extent of
Impact
from
Fishing
Gear
Current
Geographic
Extent
of Use in
Region
Overlap
of use
with coral
habitat
Overall
Rating
of Gear
Impact
Bottom Trawl N/A High High N/A N/A N/A
Mid-water Trawl N/A Low Low N/A N/A N/A
Dredge N/A High Low N/A N/A N/A
Bottom-set
Longline
N/A Med Low Low N/A N/A
Bottom-set
Gillnet
N/A Med Med N/A N/A N/A
Pelagic longline
Tuna/
swordsh
Low Med Med Low Low
Traps
Deepwater
Shrimp
Med Med Low Low Low
Hook and line
Snappers/
groupers
Low Low High Low Low
Northwestern Hawaiian Islands have recorded
seals commuting to beds of Cirripathes sp.
whip corals (100 m) where they feed on eels.
Satellite tags attached to seals indicated certain
seals spent weeks of their foraging focused at
specic subphotic locations where surveys with
submarines have revealed red and gold corals
(Parrish et al. 2002). More recent satellite tagging
of seals at the northern extent of the Northwestern
Hawaiian Islands shows similar feeding patterns
to subphotic depths (Stewart et al. 2006). Monk
seals have also been observed from the Pisces V
submersible at more than 500 m while scientists
were conducting coral surveys (A. Baco pers.
obs.). The video of the encounter shows the
seals briey looking over the submersible and
then using the light eld from the sub to look into
holes and cracks of the bottom.
VI. STRESSORS ON DEEP CORAL
COMMUNITIES
Deep coral communities within the Pacic region
may be affected by a number of natural and
anthropogenic stressors. Natural mortality has
been attributed to smothering by sediments and
by bioerosion of the substrata at the attachment
site, which leads to toppling of colonies (Grigg
1993). Detached colonies are rarely able to
reattach.
The life history attributes of deep corals makes
them highly vulnerable to habitat damage
associated with shing gear and overexploitation
in unmanaged coral sheries (Table 4.2). Many
year classes are exposed to effects at the same
time. During intensive periods of indiscriminate
shing using bottom damaging gear, decades
of accumulated coral growth can be lost (Grigg
1993).
Fishing effects
Bottom Trawling
Mobile bottom-tending gear (e.g., trawls, dredges)
are banned in the U.S. Pacic Islands Region.
From 1967 to 1975, Soviet and Japanese trawlers
shed the seamounts at the south end of the
Emperor Chain (e.g., Coco Seamount, Milwaukee
Seamount, Colahan Seamount) and some of
the seamounts at the north end of the Hawaiian
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
169
Archipelago, including seamounts that were later
designated as part of the U.S. exclusive economic
zone (e.g., Hancock Seamount, Seamount 10,
Seamount 11, Ladd Seamount) (Uchida and
Tagami 1984). The primary shing target was
Pseudopentaceros wheeleri (armorhead) and
Beryx spp. (alfonsinos)(Uchida and Tagami 1984).
In 2003, submersible dives at Seamount 11
found a large area with coral stumps and no new
colonization (A. Baco pers. obs). The destruction
was conned to a large swath within a fairly large
precious coral bed (previously unknown). It could
not be determined if this affected area was the
result of mobile bottom-tending gear used in the
early 1970s but if so, recovery clearly requires
decades.
Longline shing
Bottom longlining is not permitted in the Pacic
Islands Region. Pelagic longlining for tuna and
swordsh is permitted and is the region’s largest
shery. Longlines must be set at least 25 miles,
and in most cases 50–75 miles from emergent
parts of the Hawaiian Archipelago (WPFMC
1991). This regulation was adopted to prevent
conicts with the coastal trolling shery but it
also reduces the possibility that the gear will
affect deep corals on the slopes and seamounts
of the Hawaiian Ridge. One exception is
Cross Seamount located ~ 100 miles south of
Oahu. A popular shing site for monolament
handline shing and some longline activity, it
has accumulated numerous large fragments of
monolament line draped over the summit (F.
Parrish pers. obs). Some of these line fragments
have been seen entangled in Gerardia sp.
colonies (A. Baco pers. obs.) and other coral
trees appear to have been damaged. As this is
the only location that impacts to deep coral from
monolament shing have been documented the
assigned impact rating is “Low.”
Traps
Bottom-set traps have been used to catch lobster
and shrimp in the Hawaiian Archipelago. The
Northwestern Hawaiian Islands trap shery for
Panulirus marginatus (Hawaiian spiny lobster)
and Scyllarides squammosus (slipper lobster) is
now closed, but had always operated in waters
shallower than deep coral habitat (Polovina
1994, Dinardo and Moftt 2007). Trap shing for
the deepwater shrimp Heterocarpus laevigatus
and Heterocarpus ensifer is a small-scale pulse
shery limited to the main Hawaiian Islands that
has landed 680 metric tons since the shery’s
inception in 1984 (PIFSC IR-06-010). The shrimp
trapping overlaps the depth range of deep corals
(Ralston and Tagami 1992; Moftt and Parrish
1992), but actual impacts to deep corals have not
been documented. Shrimp have been observed
associated with hard bottom features (Moftt and
Parrish 1992) and if shers seek hard bottom to
set their traps, there is potential for damage to
deep corals. In Table 4.2, the overall gear impact
rating of shrimp trapping was classied “Low”
because of the small size and localized nature
of the shery. However if the shery expands
the potential impacts to deep corals would be
an important consideration. There are no other
recognized trap sheries operating in the U.S.
Western Pacic Islands and if any recreational
or artesianal trapping is happening, it is at a very
small scale and in shallow depths.
Other
Fishing for reef species and bottom sh typically
rely on spearing and hook and line shing.
Spearshing is largely constrained to the
shallowest depths and is unlikely to have an effect
on deep corals. Corals might be damaged by
the 3-kg bottom weight used to lower handlines
for bottom shing or might be snagged by the
attached hooks. However, visual surveys from
submersibles have inspected popular bottom
shing sites in the main and Northwestern
Hawaiian Islands for shing impacts and have
found little or no derelict gear (Kelley et al.
2006), and there are no reports of coral bycatch
(WPFMC 2005).
Effects of other human activities
Coral Harvesting
The commercial harvest of coral is the best
documented effect to black corals and precious
corals (pink, red, and gold) within the Hawaiian
Archipelago. Coral harvesting has been subject
to management under both federal and state
regulations since the 1980s. Commercial harvest
of black coral has always been selective, collected
by scuba divers using hand tools. The deeper
precious coral beds were shed initially using
nonselective tangle net dredges, but regulations
now require the use of selective methodologies
such as a submersible (Figure 4.8). Commercial
harvesting of black and precious corals has not
been reported elsewhere in the U.S. Pacic
outside of the Hawaiian Archipelago.
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170
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
Maui Divers, Inc. established the small black
coral industry in 1960 and has kept the shery
operating at varying levels continuously to the
present. Limited commercial harvest of black
coral also occurred in two other beds off Hawaii
and Kauai. State records indicate a mean annual
harvest of black coral at 1014 kg yr
-1
from 1981
to 1997 (Grigg 2001). Between 1999 and 2002
there was an increase in demand accompanied
by an increase in harvesting (Grigg 2004). This
shery often operates with fewer than three
shers so condentiality prevents disclosure
of annual data. Aggregating data into 7-year
bins showed landings increased from 1985 to
2005 with the 1999-2005 7-year bin at 22 mt
which is more than double the prior 7-year bins
(WPRFMC 2006). Much of this increase has been
attributed to improved efciency in shing due
to the availability of detailed bathymetric maps
and adoption of GPS positioning. Although 11
genera of antipatharians have been reported in
international trade, only three species (Antipathes
cf. curvata {formerly Antipathes dichotoma}
Antipathes grandis and Myriopathes ulex) have
been commercially harvested in Hawaiian waters,
with >90% of the harvested coral consisting of A.
c.f. curvata. Other black coral species known to
exist in this region are found in deeper waters
and are not considered to be of commercial grade
(Grigg 1993).
In 1965, a bed of commercial grade pink coral
was discovered at about 400 m depth on the
Milwaukee Banks in the Emperor Seamount
Chain. In 1966, Corallium secundum was also
discovered in the Makapuu Bed off Oahu, and a
small group of shermen dredged the bed using
tangle nets (Grigg 1993). Maui Divers of Hawaii
Figure 4.8. A derelict coral dredge lost during the earliest days of the shery (Dredge photo credit: A. Baco,
WHOI). Inset is the Deepworker submersible which was the most recent harvesting tool employed. Photo
credit: American Deepwater Engineering.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
171
began using a manned submersible to selectively
harvest pink, gold, and bamboo coral from this
bed. They removed a total of 8227 kg of pink
coral and 2097 kg of gold coral between 1973
and 1978 and then discontinued shing as a
result of high operating costs (Grigg 1993). Pink
corals were also harvested in 1988 from Hancock
Seamount using nonselective gear, although only
450 kg of C. secundum were obtained, most of
which were considered poor quality (Grigg 1993).
In 1978, an undescribed species of Corallium
was discovered at Midway Grounds (Emperor
Seamounts) at depths of 900–1500 m. This
resulted in a “coral rush,” with more than 100
boats from Japan and Taiwan operating in this
area. Total yield exceeded 300 metric tons from
1979 to 1984 and then dropped off because of
resource depletion (Grigg 1993).
In 1999, a Hawaii-based marine salvage and
engineering company bought two deep-worker
submersibles and began commercial harvesting
of deep corals at the Makapuu and Keahole coral
beds. Operations targeted pink, red, and gold
corals. Harvesting ended in 2001, when their
rst coral auction indicated the price of the coral
was too low to make submersible operations cost
effective, and potential harvesting grounds in the
Northwestern Hawaiian Islands were eliminated
as a result of Presidential Executive Order 13196,
which formed the Coral Reef Ecosystem Reserve
in the Northwestern Hawaiian Islands (Grigg
2002). Because the shery is made up of a single
company, condentiality prevents reporting of
landings data. However the permitted quota was
not lled at either of the two beds where corals
were harvested. Grigg (2002), working closely
with the industry, reported removal of 60% of the
allowed coral quota (1,216 kg) at the Makapuu
Bed and 20% (211 kg) at the Keahole Bed. The
precious coral shery remains dormant today.
Illegal coral dredging
Currently, the threat of illegal coral dredging is
thought to be remote. It is included here because
foreign shing vessels were documented illegally
coral dredging in the remote Northwestern
Hawaiian Islands in the early 1970s (Grigg
1993). Currently, there is no evidence or even
rumors of such illegal activity. However, much of
the Pacic region is remote and unpopulated and
any such activity could go undetected. Given the
slow growth of deep corals and low recruitment
rates, even brief periods of illegal dredging could
have lasting effects.
Invasive species
In 1972, the alien soft coral Carijoa riisei (Family
Clavulariidae) was found in the fouling community
of Pearl Harbor (Englund 2002). Originally
thought to have colonized from the tropical
Atlantic, recent genetic work (Samuel Kahng pers.
comm.) suggests it arrived from elsewhere.
It has and continues to spread to other
suitable areas in Hawaii with high ow and
low light (Figure 4.9). In 2001 deepwater
surveys of the Auau Channel black coral
beds using submarines revealed that more
than 50% of the black coral, particularly the
deeper, large reproductive colonies, were
overgrown and killed by Carijoa (Kahng and
Grigg 2005). However, Carijoa was rare on
black coral trees in waters shallower than
75 m (Boland and Parrish 2005). Light
levels are thought to be too high for Carijoa
to colonize the shallower black coral trees.
This invasive coral has been identied as a
risk to the black coral shery. Historically,
black coral trees that were too deep to be
harvested by divers were thought to serve
as a de facto reserve for the shery. With
the recent discovery that many of the deep
colonies have been killed by Carijoa, current
management practices are being reviewed
(Grigg 2004). Preliminary surveys of black
Figure 4.9. The invasive gorgonian octocoral Carijoa riisei,
that infests the deeper black corals of the Auau Channel
beds.Photo credit: F. Parrish, NOAA Fisheries.
PACIFIC ISLANDS
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
coral beds on Kauai and the island of Hawaii
have not found an infestation of Carijoa. The
occurrence of Carijoa or other invasive species
on deep corals elsewhere in the Pacic has not
been reported.
VII. MANAGEMENT OF FISHERY
RESOURCES AND HABITATS
The Pacic Islands Region has a 25-year history
of managing deep corals associated with its
Precious Coral Fishery Management Plan. The
plan was the rst shery management plan
approved by NMFS when the Western Pacic
Fishery Management Council was established
as part of the national network of shery councils
(WPFMC 1980). The plan was implemented in
September 1983 (48 FR 39229).
Management of shery resources and habitats
Historically, research has been focused on
taxonomy (Grigg and Bayer 1976), coral
assessments and estimates of age and growth
to support the information needs to manage
the shery (Grigg 1965, 1974, 1988b, 1993,
2001). In recent years, management research
has expanded to include work assessing the
connectivity of coral taxa across the Hawaiian
Archipelago (Baco and Shank 2005; Baco et al.
2006; Baco in prep.) and potential ecological
associations between corals and other fauna
(Parrish et al. 2002; Boland and Parrish 2005;
Parrish 2006; Baco and Shirley in prep.). Studies
are now focusing on the threat Carijoa riisei
presents to the black coral stock (Grigg 2002;
Kahng and Grigg 2002). Replanting corals has
been discussed as a means to mitigate impacts
to the black coral stock from Carijoa and coral
harvesting. Some preliminary “replanting”
research has been conducted with Hawaiian black
coral (Montgomery 2002), and coral harvesters
have expressed interest in continuing the work.
The expeditions that supported all this research
established study sites, deployed thermographs,
and marked colonies for future remeasurement
to validate growth and monitor the deep coral
ecosystem.
Mapping Research
Future coral research will have the benet of recent
multibeam sonar mapping efforts. Supplementing
earlier sidescan sonar (GLORIA system) and
single beam sonar mapping efforts, multibeam
products, including detailed bathymetry and
backscatter imagery, have been made for the
Hawaiian archipelago and other portions of the
U.S. Pacic (Products by John Smith at HURL;
Miller et al. 2003; Parke and Wang 2005). These
efforts will provide a fundamental bathymetric
context that future coral surveys will be able to use
to infer the likelihood of deep corals. Efforts are
currently underway to test laser-line scan survey
technology on the black coral beds of the main
Hawaiian Islands as a more promising means of
directly surveying the colony abundance of deep
corals.
Fisheries Management Council
The Western Pacic Fishery Management Council
(WPFMC) has responsibility for preparing shery
management plans (FMPs) for the sheries in the
U.S. exclusive economic zone (EEZ) of the Pacic
Islands Region. Because of the steep relief of
many Pacic Islands, deep corals also occur
within state and territory waters, and sheries
can also be governed by state and territory laws
and regulations. The Freely Associated States
are sovereign countries and management of
sheries within their EEZs is governed by their
own laws and regulations. WPFMC, an early
leader in managing habitat impacts of shing
gears, prohibited demersal sh trawls, bottom-
set longlines, and bottom-set gillnets throughout
the U.S. Pacic Island EEZ in 1983. The State
and territorial laws of Hawaii, Guam, CNMI, and
American Samoa all prohibit the use of demersal
sh trawls within their waters. The sovereign
territories of Kingman Reef, Palmyra Atoll, Jarvis
Island, Howland Island, Baker Island, Midway
Island, and Rose Atoll are National Wildlife
Refuges administered by the U.S. Fish and
Wildlife Service, while Wake Island and Johnston
Atoll are managed by the Department of Defense.
Commercial shing is not allowed within the Fish
and Wildlife Refuges. Thus, throughout the
region, both shallow and deep corals have been
largely spared impacts from trawling, at least
within the last 25 years.
Directed Harvest
The Precious Coral FMP and its regulations
classify known coral beds within the western
Pacic region and designate the harvesting
method and amount of corals that can be
harvested from each bed. All the known coral
beds are in the Hawaiian Archipelago but the
FMP includes provisions for exploratory shing
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
173
in other areas of the U.S. Pacic. The beds are
classied as: 1) Established Beds, 2) Conditional
Beds, 3) Refugia Beds, and 4) Exploratory Permit
Areas. Established beds have a history of harvest
for which maximum sustainable yield (MSY) has
been determined. Makapuu is the only designated
Established Bed although the Auau black coral
bed is in the process of being designated as
established. Conditional beds have MSYs
estimated based on their perceived size relative
to established beds. There are four conditional
beds: Keahole Point, Kaena Point, Brooks Banks
and 180 Fathom Bank. The WESTPAC Bed is
designated as a refugia bed, where no harvest
is permitted. Exploratory permit areas include
four unexplored portions of the EEZ around
Hawaii, Guam and CNMI, American Samoa,
and all remaining U.S. Island Possessions. The
FMP, as amended in 2002, prohibits the use of
nonselective gear (e.g., tangle nets, dredges)
throughout the management area. Black coral
is primarily found in State waters and the State
and the WPFMC jointly manage the resource.
Quotas and minimum size limits are monitored
through mandatory reporting to NMFS and the
Hawaii State Division of Aquatic Resources using
coral landing logs and buyer reports.
Currently, two precious coral issues are
progressing through the WPFMC process. The
rst is reconciling coral lifespan estimates derived
from radiometry studies (Roark et al. 2006) with
prior estimates made from the size structure
distribution of coral colonies and ring counts
from basal stem cross sections. Of the three
commercial corals, the black coral (Antipathes cf.
curvata) radiometric estimates were consistent
with growth rates estimated from size structure
data (Grigg and Bayer 1976). The radiometric
life span of pink coral was twice prior estimates,
and gold coral (Gerardia sp.) was estimated at
more than an order magnitude longer lived than
prior growth estimates (Grigg 2002). This has
prompted the WPFMC to put a 5-year moratorium
on the shing of gold coral until the conicting
lifespan data can be resolved. The second
issue is concern that the Maui black coral bed
may be experiencing reduced recruitment (Tony
Montgomery, State of Hawaii, unpublished data;
WPFMC 2006). This uncertainty combined with
the loss of a portion of the stock to Carijoa riisei
leaves today’s biomass at least 25% lower than
assessments in 1976 (Grigg 2004).
Closed areas
As noted above, all U.S. State and Federal
waters in the Pacic Islands are closed to
trawling and dredging—the shing techniques
most destructive to deep corals. Additional
restrictions on shing and other potentially
harmful activities are in place in the National
Wildlife Refuges, Papahānaumokuākea Marine
National Monument, and in marine protected
areas within state or territory waters.
The only area that was specically closed to
protect deep corals was WESTPAC Bank (located
N.W. of Nihoa) in the Northwestern Hawaiian
Islands. It was set aside by the precious coral
FMP as a refuge from coral harvesting. Despite
some interest, domestic precious coral shing
has never occurred in the Northwestern Hawaiian
Islands. On determining that monk seals
were visiting precious coral beds, the WPFMC
proposed expanding the refuge to include areas
where seals were visiting. Superseding this move,
the Northwestern Hawaiian Islands Coral Reef
Ecosystem Reserve was established in 2001 by
Executive Order (No. 13178 and No. 13196) and
prohibited most commercial shing, including
all harvesting of deep corals in the Reserve. In
2006, while the Northwestern Hawaiian Islands
Reserve was undergoing the designation process
to become a national marine sanctuary, it was
proclaimed a national monument by Presidential
Order under the Antiquity Act of 1906 and
renamed Papahānaumokuākea Marine National
Monument. Within the main Hawaiian Islands
and elsewhere in the Pacic, marine protected
areas have not been established specically for
the purpose of protecting deep coral communities.
However, there is interest from managers and
coral harvesters to establish a closed area off
Maui specically for black coral to serve as a
reproductive reserve and a biological reference
site.
Minerals Management Service
Oil or gas exploration does not occur in the
Pacic Islands Region. Historically, some
research has focused on the prospect of
mining manganese nodules that are formed at
abyssal depths. Recently, interest in cobalt-rich
manganese mining has resurged and large areas
of the Pacic seabed, some of which include
U.S. Pacic Islands and seamounts, are part of
the potential mining areas (International Seabed
Authority www.isa.org.jm/en/seabedarea/default.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
asp). Further studies of these cobalt-rich regions
to determine deep-coral abundance and potential
mining impacts should be considered a high
priority.
VIII. REGIONAL PRIORITIES TO
UNDERSTAND AND CONSERVE
DEEP CORAL COMMUNITIES
The priorities for future work can be divided into
the following: 1) mapping and species inventory
of deep corals, and 2) determining the important
physiological and ecological components of
deep coral ecosystems. The inherently fragile
and patchy nature of deep corals means that
determining where they are found is a primary
goal. Good success has been achieved using
available mapping, remotely operated vehicles
(ROV), and submersible infrastructure. This
should continue and extend out to the more
remote areas of the U.S. Pacic Islands Region.
Also, research is needed to validate promising
new tools to assess coral stocks with an initial
focus of using such gear at sites previously
surveyed using visual methods.
1) Highest mapping and assessment priorities
Assessments in the remote Pacic—
Historically, most research has occurred
in and around the Hawaiian Archipelago,
leaving American Samoa, Guam, CNMI, and
the rest of the western Pacic unstudied.
Baseline assessments are needed for these
other areas, particularly those that may be
affected by cobalt-manganese mining or
shery activities. Documenting areas with
extensive coral resources will permit more
focused enforcement and conservation
effort. These assessments will also provide
the DNA material for connectivity work,
provide samples to improve taxonomy and
systematics of deep corals, and provide an
invaluable test to current theories on deep
coral biogeography.
Deeper surveys—Although the Hawaiian
Archipelago has had some studies, few
baseline assessments of deep corals have
occurred outside of precious corals depth
and none below 1800 m. There is a need to
survey deeper habitats to better determine the
species ranges, biodiversity, and abundance
of deep corals.
Taxonomy—Critical to all of the assessment
and ecology studies will be a dedicated effort
to improve the taxonomy and systematics of
deep corals, and to increase the number of
people trained to identify these corals. There
are very few deep coral experts in the world
and there are currently more groups of deep
corals needing revision and new species
needing description than these taxonomists
can complete in their lifetimes.
Coral recovery studies—Beds that have
been commercially harvested or impacted by
shing gear and coral harvesting should be
periodically reassessed to determine whether
or not the coral taxa are recovering. The
seamounts that were subjected to bottom
trawling or illegal harvesting more than 30
years ago should be surveyed for signs of
coral recovery.
2) Physiological and ecological components of
the deep coral ecosystem
Environmental parameters for deep corals—
The patchy nature of deep corals, even in
areas with similar substrate, relief, and depth
implies that their distribution is inuenced
by other biological or environmental factors.
Understanding the oceanographic factors
that inuence coral distributions will be
fundamental to evaluating deep corals as a
climate record as well as predicting where
they might occur in unexplored areas.
Life history, population connectivity, and
biogeography—There is a need to understand
more about the life history, reproduction,
recruitment, growth, and dispersal abilities
of deep corals; how the populations are
connected within island/seamount chains
and between them, and how the islands and
seamounts of the U.S. Pacic are connected
genetically and biogeographically to other
parts of the Pacic.
Species associations—The ecological
contribution of corals to their associated
community needs greater attention,
particularly the invertebrates, which are more
likely to be dependent on the coral colonies.
Subphotic sh communities are likely to be
different in the remote regions of the Pacic
and may be more tightly associated with deep
corals.
The ecological impact of Carijoa riisei—
Carijoa riisei represents the most clear and
present threat to the stock of black coral and
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
175
has important implications to the black coral
shery. Research is needed to identify and
understand possible ecological changes to
the community and develop means to limit
the spread of Carijoa riisei to other black
coral habitats. Some research should be
conducted to determine the feasibility of
remediation efforts for black coral through
replanting programs.
IX. CONCLUSIONS
Deep corals occur throughout the U.S.
Pacic but only the Hawaiian Archipelago
and Line Islands have been the subject
of any surveys. Coral habitat is patchy,
suggesting at least a basic need for
suitable bottom type and conditions of rapid
ow. The gradients in dissolved oxygen,
temperature, suspended particulates, etc.,
are less understood and are a priority for
future work. Available surveys indicate
coral beds dense with colonies that cover
large areas are the exception. Given the
region has little history of trawling and
mobile bottom-tending gear, it is reasonable
to assume this is the natural condition.
Hawaiian sh are known to opportunistically
use the corals as shelter and to some
degree they co-occur with corals in high
ow habitats. It is not known how the
sh behave with deep corals in the other
parts of the Pacic. Invertebrates are
largely unstudied and the degree of their
association with deep corals is unknown
but likely to be greater.
Currently, the greatest threat to corals is the
potential for spread of the invasive species
Carijoa riisei from the Auau channel to other
black coral beds on Kauai and Hawaii.
Following that, the harvesting in the 3 beds
where the coral shery operates needs to
be closely monitored. For the black coral
bed in the Auau channel, attention is needed
because of the unanticipated loss of black
coral to Carijoa riisei. At Makapuu, the
regrowth of pink coral has been documented
once and should be checked for continued
resilience. The Keahole bed was targeted
for red and gold coral, and its prospects
for resilience are as yet unproven. Finally,
the impacts to deep corals from derelict
handlines/longlines at Cross Seamount
and shrimp trapping in the main Hawaiian
Islands should be assessed.
Many of the new coral beds that have
been identied in the Northwestern
Hawaiian Islands are protected as part
of the Papahānaumokuākea Marine
National Monument. Being remote from
the anthropogenic inuences of the main
Hawaiian Islands make them important
biological reference sites for future
research.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
183
Higher Taxon Species Distribution
Depth Range
(m) References
Phylum Cnidaria
Class Anthozoa
Subclass HExacorallia
(Zoantharia)
Order Scleractinia
Family Anthemiphylliidae Anthemiphyllia macrolobata HI Islands
369
***
Cairns, 1999
Anthemiphyllia pacica
HI Islands & Bikini Atoll,
RMI 205-296
**
Vaughan, 1907, Cairns, 1984
Family Caryophylliidae Anomocora sp. cf. A. fecunda HI Islands 201-271** Pourtalès, 1871, Cairns, 1984
Bourneotrochus stellulatus HI Islands 274-336** Cairns, 1984
Caryophyllia atlantica HI Islands 1602** Duncan, 1873, Cairns, 1984
Caryophyllia hawaiiensis HI Islands 44-388** Vaughan, 1907, Cairns, 1984
Caryophyllia marmorea HI Islands 331-337** Cairns, 1984
Caryophyllia octopali HI Islands 457-627** Vaughan, 1907, Cairns, 1984
Caryophyllia rugosa HI Islands 137-439** Moseley, 1881, Cairns, 1984
Caryophyllia sp. cf. C. ambrosia HI Islands 56-206** Alcock, 1898, Cairns, 1984
”Ceratotrochus” laxus HI Islands 583-678** Vaughan, 1907
Coenosmilia inordinata HI Islands 244-322** Cairns, 1984
Conotrochus funicolumna HI Islands 165-600** Alcock, 1902, Cairns, 1984
Crispatotrochus rubescens
HI Islands & Christmas
Island, Line Islands 197-634** Moseley, 1881, Cairns, 1984
Deltocyathus sp. cf. D. andamanicus HI Islands 274-518** Alcock, 1898, Cairns, 1984
Appendix 4.1. List of known species of ceep corals from the U.S. Pacic Islands. All species listed are found in Hawaii except one
octocoral, Keroeides koreni and several antipatharins as noted. List for octocorals and scleractinians in Hawaii based primarily on
unpublished list compiled by Dr. Stephen Cairns, Smithsonian Institution, with additions from recent Pisces cruises led by A. Baco and
additions for non-Hawaiian islands based on cited literature. List for antipatharians from Hawaii and Guam unpublished list compiled
by and courtesy of Dr. Dennis Opresko.
* = depth range known from full range for species, including outside chapter region
** = depth range known from 2 or more specimens from Hawaii, Christmas, Line Islands, etc
*** = depth from a single individual from Hawaii, usually the holotype
PACIFIC ISLANDS
184
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
Higher Taxon Species Distribution
Depth Range
(m References
Desmophyllum dianthus HI Islands Esper, 1794
Paracyathus molokensis HI Islands 161-260** Vaughan, 1907
Trochocyathus aithoseptatus HI Islands 371-454** Cairns, 1984
Trochocyathus burchae HI Islands 64*** Cairns, 1984
Trochocyathus gardineri HI Islands 274-470** Vaughan, 1907, Cairns, 1984
Trochocyathus mauiensis HI Islands 174-278** Vaughan, 1907
Trochocyathus oahensis HI Islands 75-571** Vaughan, 1907, Cairns, 1984
Trochocyathus patelliformis HI Islands
1020*** Cairns, 1999
Trochocyathus rhombocolumna HI Islands
110-530* Alcock, 1902
Family Dendrophylliidae Balanophyllia desmophyllioides HI Islands 143-406** Vaughan, 1907
Balanophyllia diomedeae HI Islands 110-307** Vaughan, 1907, Cairns, 1984
Balanophyllia gigas HI Islands
90-640* Moseley, 1881
Balanophyllia laysanensis HI Islands 238-271** Vaughan, 1907
Cladopsammia echinata HI Islands 295-470** Cairns, 1984
Cladopsammia eguchii HI Islands Wells, 1982
Eguchipsammia gaditana HI Islands 244-470** Duncan, 1873, Cairns 1984
Eguchipsammia stula HI Islands Alcock, 1902
Eguchipsammia serpentina HI Islands 269-362** Vaughan, 1907
Enallopsammia rostrata HI Islands 362-583** Pourtalès, 1878, Cairns, 1984
Endopachys grayi HI Islands 37-274**
Milne-Edwards & Haime, 1848a & b,
Cairns, 1984
Family Flabellidae Flabellum marcus
HI Islands & NW of
Wake Island 1261-1602** Keller, 1974, Cairns, 1984
Flabellum pavoninum HI Islands 183-517** Lesson, 1831, Cairns, 1984
Flabellum vaughani HI Islands 232-369** Cairns, 1984
Javania exserta HI Islands 400*** Cairns, 2006
Javania fuscus HI Islands 13-271** Vaughan, 1907
Javania insignis
HI Islands & Christmas
Island, Line Islands 52-825** Duncan, 1876, Cairns, 1984
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
185
Higher Taxon Species Distribution
Depth Range
(m References
Javania lamprotichum
HI Islands & Johnston
Atoll 244-322** Moseley, 1880, Cairns, 1984
Placotrochides minuta HI Islands 119-291*** Feinstein and Cairns, 1998, Cairns 2006
Polymyces wellsi HI Islands 440-858** Cairns, 2006
Family Fungiacyathidae Fungiacyathus ssilis HI Islands 212-503** Cairns, 1984
Fungiacyathus fragilis HI Islands 1762-2056** Sars, 1872, Cairns, 1984
Family Gardineriidae Gardineria hawaiiensis HI Islands 369-541** Vaughan, 1907, Cairns, 1984, 2006
Family Guyniidae Guynia annulata HI Islands 64-384** Duncan, 1872, Cairns, 1984
Family Micrabaciidae Letepsammia formosissima HI Islands 109-470** Moseley, 1876, Cairns, 1984
Family Pocilloporidae Madracis kauaiensis HI Islands
362-538** Vaughan 1907, Cairns 2006
Family Oculinidae Madrepora kauaiensis HI Islands 362-538** Vaughan, 1907, Cairns, 1984
Madrepora oculata
HI Islands 627-750** Cairns, 1984
Family Turbinoliidae Deltocyathoides orientalis HI Islands 439-494** Duncan, 1876, Cairns 1984, Cairns 2006
Family Stenocyathidae Stenocyathus vermiformis
HI Islands & S. Pacic
Seamounts 439** Pourtalès, 1868, Cairns, 1982,
Cairns, 1984
Order Antipatharia
Family Antipathidae Antipathes grandis HI Islands Verrill, 1928
Antipathes sp., cf. A. curvata HI Islands van Pesch, 1914
Antipathes n. sp., cf. A. dichotoma HI Islands Pallas, 1766
Antipathes sp. cf. A. abellum Guam Pallas, 1766
Antipathes sp., cf. A. spinuilosa Guam Schultze, 1896
Antipathes intermedia HI Islands 347-366*** Brook, 1889, Grigg and Opresko, 1977
Cirrhipathes anguina HI Islands 25-40*** Dana, 1846, Grigg and Opresko, 1977
Cirrhipathes contorta Guam
Cirrhipathes propinqua Guam
Cirrhipathes spiralis HI Islands Linnaeus, 1758, Grigg and Eldridge, 1975
PACIFIC ISLANDS
186
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
Higher Taxon Species Distribution
Depth Range
(m References
Stichopathes echinulata HI Islands 305-565*** Brook, 1889, Grigg and Opresko, 1977
Family Aphanipathidae Aphanipathes sarothamnoides Guam Brook, 1889
Acanthopathes undulata HI Islands 110-490**
van Pesch, 1914, Grigg and Opresko,
1977
Family Cladopathidae Trissopathes pseudotristicha
HI Islands & Channel
Islands 326-4539** Opresko, 2003
Trissopathes tetracrada HI Islands 375-425** Opresko, 2003
Family Leiopathidae Leiopathes glaberrima HI Islands Esper, 1792, Opresko, 1974
Leiopathes n. sp. HI Islands 403-471**
Prelim. ID by Opresko, Pisces Cruise
2003 &2004
Family Myriopathidae Myriopathes ulex HI Islands & Guam
Ellis and Solander, 1786, Grigg and
Eldridge, 1975
Myriopathes sp., cf. M. japonica HI Islands Brook, 1889
Cupressopathes abies Guam Linnaeus, 1758
Antipathella sp., cf. A. subpinnata HI Islands 455-460***
Ellis and Solander, 1786, Grigg and
Opresko, 1977
Family Schizopathidae Bathypathes alternata HI Islands 1195-1744** Brook, 1889, Pisces Cruise 2003
Bathypathes conferta HI Islands 380*** Brook, 1889, Grigg and Opresko, 1977
Bathypathes patula HI Islands Brook, 1889, Unpubl record at USNM
Stauropathes staurocrada
HI Islands & Johnston
Atoll
220-441, 1400-
1700** Opresko, 2002
Stauropathes sp. HI Islands 604***
Prelim. ID by Opresko, Pisces Cruise
2003
Umbellapathes helioanthes HI Islands 1205-1383** Opresko 2005
Umbellapathes, new species B HI Islands 742-744***
Prelim. ID by Opresko, Pisces cruise
2004
Dendropathes bacotaylorae HI Islands 408*** Opresko 2005
Order Zoanthidea
Zoanthid blue HI Islands 352-415** Chave and Malahoff, 1998
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
187
Higher Taxon Species Distribution
Depth Range
(m References
Zoanthid tan HI Islands 500-1910** Chave and Malahoff, 1998
Gerardia sp. HI Islands 343-577**
Ryland and Baco in prep., Chave and
Malahoff, 1998
Parazoanthus sp. 1 HI Islands 343-460** Chave and Malahoff, 1998
Parazoanthus sp. 2 HI Islands 332-1025** Chave and Malahoff, 1998
Subclass Octocorallia
Order Alcyonacea
Family Alcyoniidae Anthomastus sheri HI Islands 356-462** Bayer, 1952, Chave and Malahoff, 1998
Anthomastus (Bathyalcyon) robustus HI Islands Versluys, 1906, de Williams
Anthomastus granulosus HI Islands 20-201** Kukenthal 1910, Bayer, 1952
Inatocalyx sp. HI Islands de Williams
Family Clavulariidae Carijoa riisei Invasive, HI Islands Duch. And Mich., 1860
Clavularia grandiora HI Islands 966* Nutting, 1908, Bayer, 1952
Telestula corrugata HI Islands 914* Nutting, 1908, Bayer, 1952
Telestula spiculicola HI Islands 518-616* Nutting, 1908, Bayer, 1952
Telestula spiculicola robusta HI Islands Bayer, 1952
Family Nidaliidae Nidalia sp. HI Islands de Williams
Siphonogorgia alexanderi HI Islands 223-283* Nutting, 1908, Bayer, 1952
Siphonogorgia collaris HI Islands 144* Nutting, 1908, Bayer, 1952
Order Gorgonacea
Family Acanthogorgiidae Acanthogorgia sp. cf. A. striata Nutting, 1911 HI Islands 215-564** Grigg and Bayer, 1976
Acanthogorgia n. sp. HI Islands Muzik, 1979
Acanthogorgia sp. cf. A. paramuricata HI Islands 350-396** Stiasny, 1947, Grigg and Bayer, 1976
Acanthogorgia sp. HI Islands 1295*** Berntson et al., 2001
Cyclomuricea abellata HI Islands 71-396** Nutting, 1908, Grigg and Bayer, 1976
Muricella tenera HI Islands 237-2533* Ridley, 1884, Nutting, 1908
Family Anthothelidae Anthothela nuttingi HI Islands
340-465, 1387-
1820** Bayer, 1956, Grigg and Bayer, 1976
PACIFIC ISLANDS
188
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
Higher Taxon Species Distribution
Depth Range
(m References
Antothela n. sp. 1 HI Islands 1319*** Prelim. ID by Cairns, Pisces Cruise 2003
Antothela n. sp. 2 HI Islands 1804*** Prelim. ID by Cairns, Pisces Cruise 2003
Family Chrysogorgiidae Chrysogorgia arborescens HI Islands 722-914* Nutting, 1908
Chrysogorgia chryseis HI Islands 732***
Bayer and Stefani, 1988, Berntson et al.,
2001
Chrysogorgia delicata HI Islands 536-1463* Nutting, 1908
Chrysogorgia elegans HI Islands 433-634* Verrill, 1883, Nutting, 1908
Chrysogorgia avescens HI Islands 1688-1977* Nutting, 1908
Chrysogorgia geniculata HI Islands 146-616* Wright & Studer, 1889, Nutting, 1908
Chrysogorgia sp. cf. C. japonica HI Islands 750-1050**
Wright & Studer, 1889, Grigg and Bayer,
1976
Chrysogorgia paillosa HI Islands 704-1858*
Kinoshita, 1913, Grigg and Bayer, 1976,
Nutting, 1908
Chrysogorgia scintillans HI Islands 580-2050**
Bayer and Stefani, 1988, Chave and
Malahoff, 1998
Chrysogorgia stellata HI Islands 649-678* Nutting, 1908
Chrysogorgia sp. cf. C. stellata HI Islands
646-675, 830-
922*
Nutting, 1908, Grigg and Bayer, 1976,
Bayer and Stefani 1988
Chrysogorgia n. sp. (1/3R) HI Islands 1204*** Prelim. ID by Cairns, Pisces Cruise 2003
Chrysogorgia n. sp (2/5L) HI Islands 691-742** Prelim. ID by Cairns, Pisces Cruise 2004
Iridogorgia superba HI Islands 704-914* Nutting, 1908, Grigg and Bayer, 1976
Iridogorgia bella HI Islands 750-1925** Nutting, 1908, Chave and Malahoff, 1998
Iridogorgia. n. sp. HI Islands 1443*** Prelim. ID by Cairns, Pisces Cruise 2003
Metallogorgia melanotrichos HI Islands 183-1385* Wright and Studer, 1889, Nutting, 1908
Metallogorgia n. sp. HI Islands 1805*** Prelim. ID by Cairns, Pisces Cruise 2003
Pleurogorgia militaris HI Islands 2142* Nutting, 1908
Radicipes spiralis HI Islands 258** Nutting, 1908, Grigg and Bayer, 1976
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
189
Higher Taxon Species Distribution
Depth Range
(m References
Family Coralliidae Corallium abyssale HI Islands 1829-2403*** Bayer, 1956, Bayer unpubl ms
Corallium ducale HI Islands Bayer, 1955, Bayer unpubl ms
Corallium kishinouyei HI Islands 1145*** Bayer, 1996, Bayer unpubl ms
Corallium laauense HI Islands 365-580** Bayer, 1956, Grigg and Bayer, 1976
Corallium niveum HI Islands 232-282*** Bayer, 1956, Bayer unpubl ms
Corallium regale HI Islands 365-719** Bayer, 1956, Grigg and Bayer, 1976
Corallium secundum HI Islands 231-576**
Dana, 1846, Bayer, 1956, Grigg and
Bayer, 1976, Pisces Cruise 2003
Corallium imperiale HI Islands 1096*** Bayer, 1955, Pisces Cruise 2003
Corallium cf. secundum HI Islands Prelim. ID by Cairns, Pisces Cruise 2003
Corallium laauense x halmahera HI Islands Prelim. ID by Cairns, Pisces Cruise 2003
Corallium n. sp. HI Islands Prelim. ID by Cairns, Pisces Cruise 2003
Paracorallium tortuosum HI Islands 167-408** Bayer, 1956, Grigg and Bayer, 1976
Family Gorgoniidae Eunicella n. sp. A HI Islands 275-495** Grigg and Bayer, 1976
Family Isididae Acanella dispar HI Islands 275-445** Bayer, 1990
Acanella weberi HI Islands Nutting, 1910
Isidella trichotoma HI Islands 1920*** Bayer, 1990
Isidella sp. “5” HI Islands Muzik museum id
Isidella n. sp. (lyrate) HI Islands 1808*** Prelim. ID by Cairns, Pisces Cruise 2003
Keratoisis abellum HI Islands 346-465** Nutting, 1908, Grigg and Bayer, 1976
Keratoisis grandis HI Islands 1344-1582* Nutting, 1908
Keratoisis n. sp. HI Islands 305-565** Grigg and Bayer, 1976
Lepidisis nuda HI Islands
Wright and Studer, 1889, Grigg and Bayer
1976
Lepidisis olapa HI Islands 215-665** Muzik, 1978
Lepidisis paucispinosa HI Islands 539-631**
Wright and Studer, 1889, Nutting 1908,
Muzik, 1978
Lepidisis sp. HI Islands 1425*** Berntson et al., 2001
PACIFIC ISLANDS
190
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
Higher Taxon Species Distribution
Depth Range
(m References
Family Keroeididae Keroeides fallax HI Islands 238-245*** Bayer, 1956
Keroeides koreni Marshall Islands Wright & Studer, 1889, Bayer, 1956
Keroeides mosaica HI Islands 167-465** Bayer, 1956, Grigg and Bayer, 1976
Keroeides pallida HI Islands 146*** Hiles, 1899, Bayer, 1956
Family Paragorgiidae Paragorgia dendroides HI Islands 490-1910** Bayer, 1956, Chave and Malahoff, 1998
Paragorgia sp. cf. P. regalis Nutting, 1912 HI Islands 350-396** Grigg and Bayer, 1976
Paragorgia n. sp. HI Islands 350-396** Grigg and Bayer, 1976
Family Plexauridae Anthomuricea tenuispina HI Islands
428-531, 581-
688** Nutting, 1908, Grigg and Bayer, 1976
Anthomuricea sp. cf. A. divergens HI Islands 381-426** Kükenthal, 1919, Grigg and Bayer, 1976
Anthomuricea sp. cf. A. reticulata HI Islands 362-421** Nutting, 1910, Grigg and Bayer, 1976
Anthomuricea n. sp. A HI Islands Muzik, 1979
Bebryce brunnea HI Islands 167-396** Nutting, 1908, Grigg and Bayer, 1976
Bebryce n. sp. HI Islands Muzik, 1979
Muriceides sp. A HI Islands Muzik, 1979
Muriceides sp. B HI Islands Muzik, 1979
Filigella n. sp. Thesea n. sp. HI Islands Muzik, 1979
Muriceides tenuis HI Islands 232-362* Nutting, 1908, Muzik, 1979
Muriceides n. sp. A HI Islands Muzik, 1979
Muriceides n. sp. B HI Islands Muzik, 1979
New genus, n. sp. HI Islands Muzik, 1979
Paracis horrida HI Islands
Thomson & Henderson, 1906, Muzik,
1979
Paracis miyajimai HI Islands 362-531** Kinoshita, 1909, Grigg and Bayer, 1976
Paracis n. sp. A HI Islands Muzik, 1979
Paracis spinifera HI Islands 350-396** Nutting, 1912, Grigg and Bayer, 1976
Paramuricea HI Islandsensis HI Islands
350-396, 924-
1241** Nutting, 1908, Grigg and Bayer, 1976
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
191
Higher Taxon Species Distribution
Depth Range
(m References
Placogorgia n. sp. HI Islands Bayer unpubl ms
Placogorgia sp. n. A HI Islands Muzik, 1979
Placogorgia n. sp. B HI Islands Muzik, 1979
Placogorgia sp. cf. P. dendritica HI Islands 350-396** Nutting, 1910, Grigg and Bayer, 1976
Placogorgia sp. HI Islands 335-375** Grigg and Bayer, 1976
Placogorgia sp. cf. Ps. placoderma HI Islands 73, 182** Nutting, 1910, Grigg and Bayer, 1976
Pseudothesea sp. cf. Ps. orientalis HI Islands 147, 350-396**
Thom & Hend, 1906, Grigg and Bayer,
1976
Swiftia n. sp. 1 HI Islands 340-365** Grigg and Bayer, 1976
Swiftia n. sp. 2 HI Islands 350-396** Grigg and Bayer, 1976
Swiftia pacica HI Islands Muzik 1979
Thesea sp. cf. T. ramosa HI Islands 313-399** Nutting, ??, Grigg and Bayer, 1976
Villogorgia arbuscula HI Islands 315-412**
Wright & Studer, 1889, Grigg and Bayer,
1976
Villogorgia n. sp. 1 HI Islands 350-396** Grigg and Bayer, 1976
Villogorgia n. sp. 2 HI Islands 350-396** Grigg and Bayer, 1976
Villogorgia n. sp. A HI Islands Muzik, 1979
Villogorgia n. sp. B HI Islands Muzik, 1979
Villogorgia n. sp. C HI Islands Muzik, 1979
Family Primnoidae Callogorgia formosa HI Islands Kukenthal, 1907, Bayer, 1982
Callogorgia gilberti HI Islands 215-960** Nutting, 1908, Grigg and Bayer, 1976
Callogorgia n. sp. HI Islands 350-396** Grigg and Bayer, 1976
Calyptrophora agassizii HI Islands 781-1145** Studer, 1894, Grigg and Bayer, 1976
Calyptrophora angularis HI Islands 1207-3292* Nutting, 1908, Grigg and Bayer, 1976
Calyptrophora clarki HI Islands 12-1275 Bayer, 1951
Calyptrophora japonica HI Islands 216-432** Gray, 1866, Grigg and Bayer, 1976
Calyptrophora n. sp. HI Islands 344-454** Grigg and Bayer, 1976
PACIFIC ISLANDS
192
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
Higher Taxon Species Distribution
Depth Range
(m References
Calyptrophora wyvillei HI Islands 744-823* Wright, 1885, Nutting, 1908
Calyptrophora n. sp. 1 (lyrate) HI Islands 1078*** Prelim. ID by Cairns, Pisces Cruise 2003
Calyptrophora n. sp. 2 HI Islands 1807*** Prelim. ID by Cairns, Pisces Cruise 2003
Candidella gigantea HI Islands 1720-1815**
Wright & Studer, 1889, Pisces cruise
2003
Candidella helminthophora HI Islands 38-1820** Nutting, 1908, Grigg and Bayer, 1976
Fanellia euthyeia HI Islands Bayer and Stefani, 1989
Fanellia medialis HI Islands Bayer and Stefani, 1989
Fanellia tuberculata HI Islands Versluys, 1906, Bayer, 1982
Narella bowersi HI Islands 1344-1937*
Nutting, 1908, Berntson et al., 2001,
Grigg and Bayer, 1976
Narella dichotoma HI Islands Versluys, 1906, Bayer ms: 27
Narella sp. cf. N. megalepis HI Islands 215-564** Kinoshita, 1908, Grigg and Bayer, 1976
Narella nuttingi HI Islands 1350*** Bayer, 1997, Berntson et al., 2001
Narella ornata HI Islands 748-1007*** Bayer, 1995
Narella studeri HI Islands Versluys, 1906
Narella n. sp. 1 HI Islands 350-396** Grigg and Bayer, 1976
Narella n. sp. 2 HI Islands 353-417** Grigg and Bayer, 1976
Narella. n. sp. (unbranched) HI Islands Prelim. ID by Cairns Pisces Cruise 2003
Paracalyptrophora n. sp. HI Islands 367-398** Prelim. ID by Cairns Pisces Cruise 2004
Parastenella n. sp. HI Islands 517*** Prelim. ID by Cairns Pisces Cruise 2004
Plumarella n. sp. HI Islands 384-432** Grigg and Bayer, 1976
Thouarella (A.) biserialis HI Islands 439* Nutting, 1908, Grigg and Bayer, 1976
Thouarella (A.) regularis HI Islands 183-722* Wright and Studer, 1889, Nutting, 1908
Thouarella sp. cf. T. (T.) typica HI Islands 350-396** Kinoshita, 1907, Grigg and Bayer, 1976
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
PACIFIC ISLANDS
193
Higher Taxon Species Distribution
Depth Range
(m References
Order Pennatulacea
Family Anthoptilidae Anthoptilum murrayi HI Islands 426-2286* Kolliker, 1880, Nutting, 1908
Family Chunellidae Calibelemnon symmetricum HI Islands 196-1650** Nutting, 1908, Chave and Malahoff, 1998
Family Echinoptilidae Echinoptilum macintoshi HI Islands 225-232* Hubrecht, 1885, Nutting, 1908
Family Funiculinidae Funiculina sp. HI Islands 254-1940** Chave and Malahoff, 1998
Family Halipteridae Halipterus willemoesi HI Islands de Williams
Family Kophobelemnidae Kophobelemnon sp. (short stemmed) HI Islands Prelim. ID by Cairns, Pisces Cruise 2003
Family Pennatulidae Pennatula ava HI Islands 223-316* Nutting, 1908
Pennatula pallida HI Islands 402-530* Nutting, 1908
Pennatula pearceyi HI Islands 1033* Kolliker, 1880, Nutting, 1908
Pennatula sanguinea HI Islands 903-1033* Nutting, 1908
Family Protoptilidae Protoptilum wrighti HI Islands 523* Nutting, 1908
Protoptilum attenuatum HI Islands 925* Nutting, 1908
Protoptilum studeri HI Islands 97-421* Nutting, 1908
Family Umbellulidae Umbellula carpenteri HI Islands 1046-2056* Kolliker, 1880, Nutting 1908
Umbellula gilberti HI Islands 708-1951* Nutting, 1908
Umbellula jordani HI Islands 704-2403* Nutting, 1908
Family Virgulariidae Virgularia abies HI Islands 223* Kolliker, 1870
Virgularia molle HI Islands 1265-1280* Kolliker, 1880
Class Hydrozoa
Order Stylasterina
Family Stylasteridae Distichopora (Haplomerismos) anceps HI Islands 360-577** Cairns, 1978, 2005
Distichopora asulcata HI Islands 293-377** Cairns, 2005
Stylaster griggi HI Islands 322-583** Cairns, 2005
Stylaster infundibuliferus HI Islands 521-563** Cairns, 2005
PACIFIC ISLANDS
194
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
