Front Cover
 Front Matter
 Title Page
 Table of Contents
 Deep coral ecosystems of the United...
 State of the U.S. deep coral ecosystems...
 State of the U.S. deep coral ecosystems...
 State of the U.S. deep coral ecosystems...
 State of the U.S. deep coral ecosystems...
 State of the U.S. deep coral ecosystems...
 State of the U.S. deep coral ecosystems...
 State of the U.S. deep coral ecosystems...
 Back Matter
 Back Cover

Title: The state of deep coral ecosystems of the United States : 2007
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Title: The state of deep coral ecosystems of the United States : 2007
Physical Description: Book
Language: English
Creator: National Oceanic and Atmospheric Administration
Publisher: National Oceanic and Atmospheric Administration
Place of Publication: Silver Spring, Md.
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Table of Contents
    Front Cover
        Front Cover
    Front Matter
        Page i
    Title Page
        Page ii
        Page iii
        Page iv
    Table of Contents
        Page v
        Page vi
    Deep coral ecosystems of the United States: introduction and national overview
        Page 1
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    State of the U.S. deep coral ecosystems in the Alaska region
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    State of the U.S. deep coral ecosystems in the United States Pacific coast
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    State of the U.S. deep coral ecosystems in the western Pacific region
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    State of the U.S. deep coral ecosystems in the northeastern United States region
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    State of the U.S. deep coral ecosystems in the southeastern United States region
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    State of the U.S. deep coral ecosystems in the northern Gulf of Mexico region
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    State of the U.S. deep coral ecosystems in the United States Caribbean region
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    Back Matter
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    Back Cover
        Page 367
Full Text





. 4


'r 1b


Citation for the entire document:

Lumsden SE, Hourigan TF, Bruckner AW, Dorr G (eds.) 2007. The State of Deep Coral Ecosystems
of the United States. NOAA Technical Memorandum CRCP-3. Silver Spring MD

Citation for an individual chapter (e.g., Alaska Chapter):

Stone RP and Shotwell SK 2007. State of Deep Coral Ecosystems in the Alaska Region: Gulf of
Alaska, Bering Sea and the Aleutian Islands. pp. 65-108. In: SE Lumsden, Hourigan TF, Bruckner AW
and Dorr G (eds.) The State of Deep Coral Ecosystems of the United States. NOAA Technical Memo-
randum CRCP-3. Silver Spring MD 365 pp.

Cover illustration courtesy of Michael Peccini, NOAA

For more information:

For more information about this report or to request a copy, please contact NOAA's Coral Reef
Conservation Program, 301-713-0299. NOAA/NMFS/OHC 1315 East West Highway, Silver Spring,
Maryland 20910. Or visit http://coralreef.noaa.gov/


This publication does not constitute an endorsement of any commercial product or intend to be an opinion
beyond scientific or other results obtained by the National Oceanic and Atmospheric Administration
(NOAA). No reference shall be made to NOAA, or this publication furnished by NOAA, in any advertising
or sales promotion which would indicate or imply that NOAA recommends or endorses an proprietary
product mentioned herein, or which has as its purpose an interest to cause directly or indirectly the
advertised product to be used or purchased because of this publication.

NOAA Technical Memorandum CRCP 3

United States Department Nationa
of Commerce Atmosphel

Carlos M. Gutierrez Conrad C.
Secretary Adr

)ceanic and National Marine Fisheri(
Administration Service

utenbacher, Jr. William T. Hogarth
listrator Assistant Administratol


IIl Report and Chapter 1: Introduction and (
litors would like to thank Michael Peccini for devel
raphs and images for use in this chapter. We wo
isiderable time and effort to provide constructive c
,view of the full report was conducted by the Cent(
Reviewers for the CIE review included Stephen (
ips Institution of Oceanography, and Pal B. Morter
port also benefited from additional external review
stern Pacific Fishery Management Council, New E
C University of Connecticut, Andrew Shepard NI
- Hawaii Undersea Research Laboratory, Robert'
lity, John K. Reed Harbor Branch Oceanographii
4&M, John Warrenchuck and Santi Roberts Oce
Biology Institute, Alberto Lindner- Smithsonian In
il NOAA reviewers included: NOS comments from
Hickerson FGBNMS, G.P. Schmahl FGBNMS
nd Brad Barr NMSP, Jeff Hyland NCCOS, OAF
southeast Fisheries Science Center, Southeast Re(
rest Regional Office, Southwest Fisheries Science
a of Protected Resources, Robert Brock Office c
; a collaborative effort with the authors from each <
:hened this section.

:er 2: Alaska Chapter
iifetz (AFSC) provided the overview on multi-bearr
ier draft of this chapter. Jana DaSilva Lage and R
nation regarding submarine telecommunication cable
s Hole Oceanographic Institution), Peter Etnoyer (
sity) provided information on coral distribution frorr
)nservation Biology Institute) provided helpful insic
Pacific corals. Jennifer Reynolds (University of Ale
geology of the North Pacific Ocean. Cathy Coon
osures in Alaskan waters and Jon Warrenchuk (0

:er 3: West Coast
ous agencies, institutions and individuals provide
i, Southwest Fisheries Science Center), Brian Tiss
), Milton Love (Univ. California, Santa Barbara), E5
National Marine Sanctuary), Jeff Hyland (NOS, Nc
rly at Washington State Univ. Vancouver), Dan Ho
Sanctuary), Erica Burton (NMSP, Monterey Bay 1`
Fisheries Science Center), Lance Morgan (Marini
iautix Consulting), Chris Goldfinger, Chris Romsos
Landing Marine Laboratories), Rikk Kvitek (Califor
vith Duke Univ.) and Glen Jamieson (Department

tion to reviews by the Center for Independent Exp
provided by Waldo Wakefield and Ewann Berntson
ich (NMFS, Southwest Fisheries Science Center),
National Marine Sanctuary), Jeff Hyland (NOS, NE


ng GIS images and numerous others who provide
also like to thank the many reviewers for contribu
iments on this chapter. An independent external
or Independent Experts (CIE) at the University of
rns Smithsonian Institution, J. Anthony Koslow
n Institute of Marine Research, Bergen, Norway.
y the South Atlantic Fishery Management Coun-
iland Fishery Management Council, Peter Auster
C University of North Carolina, Wilmington, Chris
3eorge --George Institute for Biodiversity and Sus-
istitution, Peter Etnoyer Harte Research Institute
a, Lance Morgan and Fan Tsao Marine Conser-
:ution, Brian Tissot Washington State University.
I Bowlby OCNMS, Mary Sue Brancato --OCNMS
oger Griffis and Kara Meckley --CRCP, Steve Git-
,omments from NURP and OE, NMFS comments
nal Office, Northeast Fisheries Science Center,
enter, Alaska Regional Office, Dwayne Meadows
science and Technology. The Introductory Chap-
he regional chapters, and their input has greatly

lapping efforts in Alaskan waters and reviewed
Hansen (Fugro Pelagos, Inc.) provided helpful
deployments in Alaskan waters. Amy Baco-Taylor
ie Gulf of Alaska seamounts. John Guinotte (Ma-
; regarding the effects of ocean acidification on
a Fairbanks) provided information about the sub-
PFMC) provided detailed information on fishing
ana) provided Figure 2.13.

ata and input to Chapter 3, including Mary Yoklavic
and Jennifer Bright (Washington State Univ. Van-
lowlby and Mary Sue Brancato (NMSP, Olympic
nal Centers for Coastal Ocean Science), Jodi Pirtl
rd and Dale Roberts (NMSP, Cordell Bank Nation,
ional Marine Sanctuary), Mark Wilkins (NMFS,
conservationn Biology Institute), Peter Etnoyer
id Mark Hixon (Oregon State Univ.), Gary Greene
i State Univ. Monterey Bay), Alberto Lindner (for-
Fisheries and Oceans Canada).

s. additional reviews of all or Dortions of ChaDter 3

in and Fan Tsao (Marine Conservation Biology Ins

chapter 4: Hawaii
uch of the research reported in chapter 4: the Wes
as supported by the NOAA Office of Ocean Explor
e Hawaii Undersea Research Laboratory. Stephe
cations and unpublished species lists to help us p
'e are also grateful to Celeste Mosher, Deborah Ye

chapter 5: Northeast
ie authors thank Beth Lumsden, Tom Hourigan, ar
is effort and for editing and formatting our chapter.
airns, Pal Mortensen, and J. Anthony Koslow, as v

chapter 6: Southeast
'e thank the NOAA Office of Ocean Exploration, th
management Council (SAFMC), Environmental Defe
nd our deep coral research, which contributed to tl
team effort involving the authors, K.J. Sulak (USG
;on (USGS) and A. Howard. Andy Shepard (Natio
ted several of our projects, including this report, ar
e efforts of the SAFMC in leading the way toward
uattrin and M.L. Partyka for help with figures and c
DAA Ecosystem Assessment Division.

chapter 7: Gulf of Mexico
ie authors would like to extend their appreciation c
ive greatly improved this chapter. A special 'Thank
I History) who helped us navigate the complexities
banks National Marine Sanctuary who generously s

chapter 8: Caribbean

ite). We appreciate all their constructive comment

-n Pacific Region: Hawaii and the US Pacific Islan(
on and the NOAA Undersea Research Program th
,airns and Dennis Opresko provided preliminary ic
'ide as complete a taxonomic inventory as possible
aguchi and Ronald Hoeke who helped with tables

other members of the Deep Coral Team for leading
he authors would also like to thank reviewers Ster
as additional reviewers, for constructive comment

J.S. Geological Survey, the South Atlantic Fishery
se, and the Minerals Management Service for help
review. Much of these data were collected as par
E. Baird (NC Museum of Natural Sciences), C. M
Undersea Research Center, UNC-Wilmington) fai
provided the Oculina photographs. We acknowlec
:ter management of deep coral habitats. We thank
a analysis. This chapter was partially supported b

I thanks for all the contributions from reviewers, wl
)u' goes to Dr Steven Cairns (National Museum of
:coral taxonomy, and to the staff of the Flower Gal
red their research with us'

*nce, ana comments were invaluaDi

Ecosystems of ti

NOAA Technical Men

Table of C


of Contents


er 1: Deep Coral Ecosystems of the United
Thomas F Hourigan, S. Elizabeth Lurr
Sandra Brooke, Robert P Stone

er 2: State of the U.S. Deep Coral Ecosyste
Sea and the Aleutian Islands
Robert P Stone and S. Kalei Shotwell

er 3: State of the U.S. Deep Coral Ecosyste
to Washington
Curt E. Whitmire and M. Elizabeth Cla

er 4: State of the U.S. Deep Coral Ecosyste
United States Pacific Islands
Frank A. Parrish and Amy R. Baco

er 5: State of the U.S. Deep Coral Ecosyste
Maine to Cape Hatteras
David B. Packer, Deirdre Boelke, Vince

er 6: State of the U.S. Deep Coral Ecosyste
Cape Hatteras to the Florida Straits
Steve W Ross and Martha S. Nizinski

er 7: State of the U.S. Deep Coral Ecosyste
Straits to Texas
Sandra Brooke and William W Schroe

er 8: State of the U.S. Deep Coral Ecosyste
Puerto Rico and U.S. Virgin Islands
Steven J. Lutz and Robert N. Ginsburc

SUnited States

randum CRCP-3





rates: Introduction and National Overview
ten, Gabrielle Dorr, Andrew W Bruckner,

s in the Alaska Region: Gulf of Alaska, Bering


s in the United States Pacific Coast: California

? 109

s in the Western Pacific Region: Hawaii and t\


s in the Northeastern United States Region:

7uida, and Leslie-Ann McGee 195

s in the Southeastern United States Region:


s in the Northern Gulf of Mexico Region: Floric

.r 271

s in the United States Caribbean Region:


lis report represents the first effort by the Natii
partnership with other federal, academic and
formation on the abundance and distribution (
apths greater than 50 m. It consists of an introc
ascribing deep coral communities in U.S. water
sular Pacific, the Northeastern U.S., Southea
port reflects the tremendous increase in awar
st few years as the result of increasing explore
:eans. In the U.S., NOAA is proud to serve as

OAA coordinated the development of this repi
OAA Coral Reef Conservation Program. It reflex
gion and these teams should be cited as prin
mnefited from the comments and suggestions c
quality Act peer review coordinated through the

i introductory chapter defines and provides ba
id identifies major threats that they face. A
)mmunities across the regions from a national p
ere developed by authors considered experts
id those chapters represent the core of this re
gion and geological and oceanographic featL
ajor deep coral taxa that structure habitats in
ovide information on the other species assoc
habitats; discuss management efforts developed
formation needs.. The report also includes unr
search expeditions.

iis report fulfills a commitment made in the I
search, survey and protect deep coral comrr
e importance of these communities as hot-s
)mmitment to ensuring their enhanced conser
OAA effort to develop a National Deep Coral a
:rategy. We hope that this first Report on the
ill stimulate additional research, surveys and
)cument both increased understanding and pr

al Oceanic and Atmospheric Administration (N
n-governmental partners, to bring together avw
structure-forming corals that occur in U.S. wal
:tion, National Overview and seven regional ch
ff Alaska, the U.S. West Coast, Hawai'i and th
3rn U.S., Gulf of Mexico, and U.S. Caribbean
ess of these communities that has evolved o\
)n and research to understand deeper regions
leading partner in much of this work.

, under the auspices of the Deep Coral Team
s the work and dedication of writing teams fron
-y authors of the regional chapters. The repo
numerous federal and external reviewers and
enter for Independent Experts.

ground information on structure-forming deep
national Overview explores general trends in
spective. Chapters 2 through 8, the regional chE
1 the field of deep coral research and manag
)rt. The authors of each chapter briefly descri
s important to deep coral communities; ident
e region and what is known about their district
:ed with coral habitat; describe the threats to
i respond to these threats, and briefly outline re
)lished data and observations collected during

3. Ocean Action Plan as part of an overall el
cities. It reflects NOAA's growing understand
)ts for deep-water biological diversity, and N
ition. This report is also a central part of a br
I Sponge Research, Conservation and Manag
ate of Deep Coral Ecosystems of the United
protection, and hope that periodic future report
actionn of these unique and valuable ecosysterr



Thomas F. Hourigan1, S. Elizabeth Lumsden1, Gabrielle Dorr2, Andrew W. Bruckner1,
Sandra Brooke3, Robert P. Stone4


Coral reefs are among the most spectacular
ecosystems on the planet, supporting such rich
biodiversity and high density of marine life that
they have been referred to as the "rainforests
of the sea." These ecosystems are usually
associated with warm shallow tropical seas,
generally within recreational diving depths (30
m or less). However other coral communities

thrive on continental shelves and slopes around
the world, sometimes thousands of meters
below the ocean's surface. These communities
are structured by deep corals, also referred to
as "deep-sea corals" or "cold-water corals," and
are found in all the world's oceans. Unlike the
well-studied shallow-water tropical corals, these
corals inhabit deeper waters on continental
shelves, slopes, canyons, and seamounts in
waters ranging from 50 m to over 3,000 m in

Figure 1.1 An Alaskan "coral garden" with several
species of soft corals, hydrocorals, hydroids, and
demosponges. Photo credit: Alberto Lindner

depth. A few species also extend into shallower,
cold waters in the northern latitudes (Figure 1.1).

Deep coral habitats appear to be much more
extensive and important than previously known,
particularly with respect to supporting biologically
diverse faunal assemblages (Wilkinson 2004;
Roberts et al. 2006). At the same time, they are

1NOAA National Marine Fisheries Service,
Office of Habitat Conservation
1315 East West Hwy Silver Spring, MD 20910
2NOAA National Marine Fisheries Service,
Southwest Regional Office
3Ocean Research and Conservation Association,
Fort Pierce, Florida 34949
4Auke Bay Laboratory, National Marine
Fisheries Service, Alaska Fisheries Science Center,
11305 Glacier Highway, Juneau, Alaska 99801-8626


increasingly threatened by a variety of activities
ranging from bottom fishing to energy exploration
(Rogers 1999; Koslow et al. 2000). Over the
past decade, science has demonstrated that
deep corals are often extremely long-lived, slow-
growing animals, characteristics that make them
particularly vulnerable to physical disturbance,
especially from activities such as bottom
trawling. Where water, current, and substrate
conditions are suitable, these corals can form
highly complex reef-like structures, thickets or
groves, and there is increasing evidence that
many areas of deep coral and sponge habitats
function as ecologically important habitats for
fish and invertebrates.


Structure-forming deep corals, as used in this
report (Box 1.1), include a number of very different
species that contribute to three-dimensionally
complex habitats in deeper waters. Structure-
forming deep corals are defined as those coral
species with complex branching morphology and
sufficient size to provide substrate or refuge for
associated fishes and invertebrates. As such,
they represent a functional group of conservation
interest, rather than a taxonomic group, which
Morgan et al. (2006) have likened to the diverse
plants included under the descriptors "bushes"
or "trees." These coral species are found within
two cnidarian Classes, Anthozoa and Hydrozoa
(Box 1.2). Anthozoa includes the stony corals,

black corals, and gorgonians among the
more prominent deep coral groups, while the
Class Hydrozoa contains the stylasterid corals
(often referred to as lace corals) in the order
Anthoathecatae. As a group, deep corals are
among the most incompletely understood corals,
and field and laboratory investigations are

Deep corals in this report are distinguished
from "shallow" tropical corals, the subject of a
separate NOAA status report (Waddell 2005),
by restricting consideration to azooxanthellate
corals, meaning they lack the symbiotic algae
(zooxanthellae) found in most shallow corals

and do not require sunlight to grow. The depth
range defining "deep" corals for the purposes of
this report (>50 m), while somewhat arbitrary, is
based on the best scientific information available
(e.g., depths at which azooxanthellate corals
predominate over zooxanthellate corals) as well
as by practical conservation considerations.
Generally, "deep-sea organisms" are defined as
those occurring deeper than the continental shelf
(generally around 200 m). However, a number
of coral communities of management interest
occur at shallower depths (e.g., Oculina coral
banks off Florida and black coral beds in Hawaii),
and share functional similarities to true deep-sea
coral taxa. Even though several of these coral
species have been harvested for jewelry since
antiquity, and their existence has been known to
science since 1758 (when Carl von Linne wrote

Box 1.1 "Structure-forming deep corals" and "deep coral communities" defined:
For the purposes of this report:
Structure-forming deep corals are any colonial, azooxanthellate corals generally oc-
curring at depths below 50 m that provide vertical structure above the seafloor that can
be utilized by other species. It includes both branching stony corals that form a structural
framework (e.g., reefs) as well as individual branching coral colonies, such as gorgonians
and other octocorals, black corals, gold corals, and lace corals. Though these are often
referred to as habitat-forming deep-sea, deep-water, or cold-water corals, the more neutra
term "structure-forming" has been used in this document to avoid an implication of habitat
associations with other species until such associations have been demonstrated by the
best available science. Tables of important structure-forming coral species within the U.S.
EEZ are listed in Appendix 1.1 and 1.2.
Deep coral communities are defined as assemblages of structure-forming deep corals
and other associated species, such as sedentary and motile invertebrates and demersal


the Systema Natura) relatively little is known
about their biology, population status, the role
they play in enhancing local species diversity,
or their importance as habitat for deep-water
fishes, including those targeted by fishermen.
With recent advances in deep-sea technology,
scientists are now beginning to locate and map
the distribution of deep coral habitat, and the past
15 years has seen a rapid expansion of studies
on these deep-sea communities worldwide.

Deep corals include both reef-building and non-
reef-building corals. Although only a few stony
coral species (order Scleractinia) form deep-
water structures such as bioherms, coral banks
or lithoherms (Box 1.3) (Freiwald et al. 2004;

George 2004a, b; Cairns in press), these species
can occur as individual small colonies less than a
meter in diameter or they may form aggregations
that can create vast reef complexes tens of
kilometers across and tens of meters in height
over time (Freiwald et al. 2004; Roberts et al.

Shallow corals need well-known and well-
documented environmental conditions for
development; however the requirements for deep
coral species are not as well understood. Table
1.1 highlights some of the general differences
and similarities between shallow and deep stony
corals. The major structure-forming coral taxa
are described in a later section. Unlike stony

Box 1.2. Taxonomy of Major Coral Groups1
"Coral" is a general term used to describe several different groups of animals in the Phylum
Cnidaria. The following is a summary of cnidarian taxonomy as used in this report, showing
the primary groups containing animals referred to as "corals." Orders in bold contain structure-
forming deep corals.
I. Class Anthozoa corals, sea anemones, sea pens
I.A. Subclass Hexacorallia (Zoantharia) sea anemones, stony and black corals
I.A.1. Order Scleractinia stony corals (The most important families
containing deep-water structure-forming stony corals are
Carophylliidae, Dendrophylliidae, and Oculinidae)
I.A.2. Order Zoanthidea zoanthids (family Gerardiidae)
I.A.3. Order Antipatharia2 black corals
I.B. Subclass Octocorallia (Alcyonaria) octocorals
I.B.1 Order Alcyonacea -true soft corals, stoloniferans3
I.B.2 Order Gorgonacea4 sea fans, sea whips (there are at least 12
families containing deep-water structure-forming gorgonians)
I.B.3 Order Pennatulacea sea pens
I.B.4 Order Helioporacea Lithotelestids and blue corals
II. Class Hydrozoa hydroids and hydromedusae
II.A.1. Order Anthoathecatae5 stylasterid corals and fire corals
suborder Filifera (Stylasteridae: stylasterids, lace corals)
II. Class Cubozoa does not contain corals
IV. Class Scyphozoa does not contain corals
1Taxonomic summary generally follows that presented in the Integrated Taxonomic Information
System (http://www.itis.gov).
2 Black corals were formerly placed in the subclass Ceriantipatharia; however, based on recent
molecular data they are now considered to be in the same subclass as other hexacorals.
3 Current taxonomy has the order Stolonifera combined with Alcyonacea (S. Cairns pers. comm.)
4Not all taxonomists recognize the order Gorgonacea as separate from Alcyonacea.
SThe order containing lace corals (family Stylasteridae) was previously called Filifera or
Stylasterina. Filifera is now considered a suborder and Stylasterina is no longer valid (S. Cairns
pers. comm.).


Table 1.1 Differences between tropical shallow-water and deep-water structure-forming stony corals

0-100 m 39-3,000 m
18-310 C 4-13o C
Tropical and subtropical seas from Potentially global, at least
30oN-300S 560 S-710 N
Yes No (Note: several species of Oculina
and Madracis have a facultative
relationship with zooxanthellae in
shallow populations)
1-10 mm per year for massive slow 1-20 mm per year for Oculina and
growing corals Lophelia3; growth rates of other taxa
50-150 mm per year for faster are unknown.
growing branching corals
Approximately 800 Approximately 6-14

Photosynthesis, zooplankton and Zooplankton and possibly suspended
suspended organic matter organic matter
Overfishing and destructive fishing Bottom-tending fishing gear
Pollution and siltation Oil and gas exploration and
Coastal development Pipelines and cables
Harvest of corals Climate change (ocean acidification
and possible changes in currents and
Recreational misuse
Climate change (coral bleaching,
ocean acidification and storm
1. Modified from Freiwald et al. 2004
U.S. Coral Reef Task Force 2000 Threats to shallow coral reefs
3. Mortensen and Rapp (1998) reported rates of 25 mm/yr but this is thought to be an overestimate due to the
sampling methodology.

Box 1.3 Geological Terms (see Chapter 8 for more detail)

Bioherm A moundlike or reeflike formation built by organisms such as corals,
algae, foraminfera, mollusks, etc., composed almost exclusively of their
calcareous remains and trapped sediments, and surrounded by rock of different
physical characteristics. It may take the form of an unconsolidated coral mound
or reef, or be covered by crust-like layers of limestone (Lithoherm).

Coral bank An undersea mound or ridge that rises above the surrounding
continental shelf or slope and is formed in part from the carbonate
skeletons of corals.

Lithoherm A deep-water mound of limestone, usually formed by submarine
consolidation of carbonate mud, sand and skeletal debris


Is, other deep coral taxa, such as stylasterids, bi
onians, and black corals do not form reefs, e:
)ften have complex, branching morphologies be
may form dense groves or thickets. Sea oi
may exist either singly on the seafloor or a:
n large and complex ecosystems. The North B
fic, for example, is known to have extensive in
I "gardens" composed of gorgonians and gl
erous other coral and sponge species. lil
the understanding of deep coral is
munities and ecosystems has increased,
as appreciation of their value. Deep coral D
munities can be hot-spots of biodiversity id
e deeper ocean, making them of particular in
servation interest. Stony coral "reefs" as well vw
tickets of gorgonian corals, black corals, and al
ocorals are often associated with a large fe
ber of other species. Through quantitative w
eys of the macroinvertebrate fauna, (2
d (2002b) found over 20,000 individual la
rtebrates from more than 300 species living al
ng the branches of ivory tree coral (Oculina cc
;osa) off the coast of Florida. Over 1,300 (
;ies of invertebrates have been recorded in A
ongoing census of numerous Lophelia reefs ol
ie northeast Atlantic (Freiwald et al. 2004), re
Mortensen and Fossa (2006) reported 361 st
;ies in 24 samples from Lophelia reefs off be
vay. Gorgonian corals in the northwest vw
itic have been shown to host more than al
species of invertebrates (Buhl-Mortensen al
Mortensen 2005). An investigation by oi
er de Forges et al. (2000) reported over 850 h,
ro- and megafaunal species associated with gi
nounts in the Tasman and south Coral Seas A
many of these species associated with Ir
deep coral Solenosmilia variabilis (Rogers cc
1). The three-dimensional structure of deep re
Is may function in very similar ways to al
tropical counterparts, providing enhanced
ing opportunities for aggregating species, a D
ig place from predators, a nursery area for tlh
niles, fish spawning aggregation sites, and c.
:hment substrate for sedentary invertebrates a
sa et al. 2002; Mortensen 2000; Reed cl
?b). a

)gical research on marine organisms. For
nple, several deep-water sponges have
i shown to contain bioactive compounds
pharmaceuticall interest; sponges are often
)ciated with deep coral communities.
iboo corals (family Isididae) are being
stigated for their medical potential as bone
ts and for the properties of their collagen-
gorgonin (Ehrlich et al. 2006). A number of
) corals are also of commercial importance,
ecially black corals (order Antipatharia) and
and red corals (Corallium spp.), which are
Dasis of a large jewelry industry. Black coral
awaii's "State Gem."

p coral communities have also been
tified as habitat for certain commercially-
)rtant fishes. For example, commercially
able species of rockfish, shrimp, and crabs
known to use coral branches for suspension
ing or protection from predators in Alaskan
irs (Krieger and Wing 2002). Husebo et al.
12) documented a higher abundance and
er size of commercially valuable redfish, ling,
tusk in Norwegian waters in coral habitats
pared to non-coral habitats. Costello et al.
15), working at several sites in the Northeast
itic, report that 92% of fish species, and 80%
idividual fish were associated with Lophelia
habitats rather than on the surrounding
)ed. Koenig (2001) found a relationship
/een the abundance of economically
able fish (e.g., grouper, snapper, sea bass,
amberjack) and the condition (dead, sparse
intact) of Oculina colonies. Oculina reefs
-lorida have been identified as essential fish
tat for federally-managed species, as have
Ionian-dominated deep coral communities off
ka and the West Coast of the United States.
their cases, however, the linkages between
mercial fisheries species and deep corals
ain unclear (Auster 2005; Tissot et al. 2006)
may be indirect.

to their worldwide distribution and the fact
some gorgonian and stony coral species
live for centuries, deep corals may serve as
oxy for reconstructing past changes in global
ate and oceanographic conditions (Risk et
1002; Williams et al. 2007). The calcium
onate skeletons of corals incorporate trace
ients and isotopes that reflect the physical
chemical conditions in which they grew.
lysis of the coral's microchemistry has


allowed researchers to reconstruct past oceanic


The term "coral" is broadly used to describe a
polyphyletic assemblage of several different
groups of animals in the phylum Cnidaria and
includes a range of taxa (Box 1.2). Structure-
forming corals outlined in this document are
animals in the cnidarian Classes Anthozoa
and Hydrozoa that produce calcium carbonate
aragonitee or calcite) secretions. These
secretions have different forms: a continuous
skeleton, numerous, usually microscopic,
individual sclerites, or a black, horn-like,
proteinaceous axis (Cairns in press). The
following are the major classes and orders that
include important structure-forming deep corals.
Species identified in this report as important
structure-forming corals in U.S. waters are
shown in Appendix 1.1 and 1.2.


Anthozoa, the largest Class of cnidarians,
contains over 6,000 described species (Barnes
1987). They are found as both solitary and
colonial arrangements. They have a cylindrical
body shape with an oral opening surrounded
by tentacles, and have lost the medusoid
(medusa or jellyfish shape) life history stage.
In anthozoans, the mouth leads through the
pharynx to the gastrovascular cavity, a feature
unique to cnidarians that serves both a digestive
and a circulatory function. This cavity is divided
into compartments radiating outward from the
pharynx and is lined with nematocysts.



Stony corals (order Scleractinia) are exclusively
marine anthozoans with over 1,400 described
species. Individual polyps secrete a rigid external
skeleton composed of calcium carbonate in
the crystal form aragonite. Over 776 of the

recognized stony corals are found in shallow
water and contain zooxanthellae (symbiotic
algae) that provide much of the coral's nutrition,
while deep-water species lack zooxanthellae.
While more than 90% of the shallow stony
corals are colonial structure-forming species
(many contributing to coral reefs), there are at
most 14 species of azooxanthellate deep-water
scleractinians in the world that can be considered
structure-forming species, 13 of which occur in
U.S. waters (Cairns 2001; Cairns in press). The
other 97.7% of the deep-water species are for
the most part small (some as small as 2 mm
adult size) and solitary (74%) (Cairns 2001).
Two deep corals that are major contributors to
reef-like structures or bioherms in U.S. waters
(Lophelia pertusa, and Oculina varicosa) while
other stony corals including Madrepora oculata,
Solenosmilia variabilis, and Enallopsammia
profunda contribute to the formation of bioherms
and reefs in some areas. Goniocorella dumosa
(Alcock 1902) is an important framework-
building coral found in the southwest Pacific
Ocean, especially around New Zealand, where
it can form large, localized reefs up to 40 m in
height and 700 m wide. G. dumosa appears to
be restricted to the southern hemisphere, and
has not been reported from U.S. waters (Cairns


I.A. 1.a.i. Lophelia pertusa
(Linnaeus, 1758)1

Description: Lophelia pertusa belongs to the
family Caryophylliidae, Vaughan and Wells, 1943.
At present the genus Lophelia is monotypic
(Zibrowius 1980). A number of different Lophelia
species were described previously, but were
either synonymous with L. pertusa or reclassified
into other genera (for a list of synonyms see
Rogers 1999). Worldwide, L. pertusa is the
most important constituent of deep-water coral
reefs, forming massive complexes hundreds of
kilometers long and up to 30 m high (Freiwald et
al. 2004). L. pertusa is often found in association
with E. profunda, M. oculata, and S. variabilis in
1Note on nomenclature: The name of the author
who described the species follows the species
name, e.g., Solenosmilia variabilis Duncan, 1873. If
subsequent work has placed a species in a different
genus, the author's name appears in parentheses,
e.g., Enallopsammia profunda (Pourtales, 1867).


1-igure 1.z samples OT Lopenla pertusa colonies collectea Trom tne Uult oT Mvexico. I ne lent
specimen displays the more heavily calcified "brachycephala" morphology with large polyps, and
the right specimen shows the more fragile graciliss" morphology. Photo credit: Sandra Brooke,
OIMB, Charleston, OR.

the western Atlantic, along the Blake Plateau, and
along the Florida-Hatteras slope (Reed 2002b).
Lophelia is fragile, slow growing, and extremely
susceptible to physical destruction from fishery
impacts (Fossa et al. 2002; Reed 2002b).

Distribution: L. pertusa is a widespread
structure-forming deep-water scleractinian
species occurring in the Atlantic, Pacific, Indian,
and Southern Oceans, with a latitudinal range
from about 560 S to 710 N (Freiwald et al. 2004).
In U.S. waters major reefs have been reported
off the southeastern U.S. (Chapter 6) and the
Gulf of Mexico (Chapter 7). The species has also
been reported from the West Coast (Chapter 3),
the Caribbean (Chapter 8) and the New England
Seamounts (Chapter 5).

Depth Range: L. pertusa has been recorded
from depths as shallow as 39 m in the Norwegian
fjords (Freiwald et al. 2004) and as deep as 2,170
m (Cairns 1979), but most commonly forms reefs
at depths between 200 m and 1,000 m (Freiwald
et al. 2004).

Morphology: This species displays great
phenotypic plasticity in colony morphology
ranging from heavily calcified structures with
large polyps (1-1.5 cm in diameter) termed
"brachycephala" by earlier workers, to the more
delicate graciliss" morphology with smaller
polyps and more defined septal ridges (Figure
1.2; Newton et al. 1987). Lophelia colonies can
exhibit great morphological variation, which may
reflect the local environmental conditions of their
habitat, but characteristically form bushy thicket-
like structures composed of living branches
overlying a center of dead coral (Figure 1.3).
Branches are dendritic and readily fuse together,
which increases colony strength.

Growth and Age: The growth rate of L. pertusa
in the northeast Atlantic has been estimated
at 5-26 mm yr1 (Mortensen and Rapp 1998;
Mortensen 2001; Gass and Roberts 2006),
suggesting that large colonies probably represent
hundreds of years of accretion. Radioisotope
dating of Lophelia reefs from seamounts off
northwest Africa, the Mid-Atlantic Ridge, and
the Mediterranean suggest that they may have


Figure 1.3 Colonies of living and dead Lophe

grown continuously for the last 50,000 y,
(Schroder-Ritzrau et al. 2005).

Reproduction: L. pertusa is a gonochor
species (separate sexes) that produces a si
, .

cohort of about 3,000 relatively small (mz
140 pm in diameter) oocytes per polyp E
year (Waller and Tyler 2005). The species i

Figure 1.4 Sesiia ariabis coral. Photo
Figure 1.3 Colonies of smilia variabilis coral. Photo

with squat lobster. Photo credit: Ross et al.,

*s annual broadcast spawner, releasing gami
between January and February (Le Goff-)
and Rogers 2005; Waller and Tyler 20
ic The low genetic diversity in some locatic
e the occurrence of genetically distinct 1
= and offshore populations, and the prese
:h of lecithotrophic larvae suggest there is a I
n degree of local recruitment (Le Goff-Vitry
Rogers 2005). Local recruitment, toge
with predominance of asexual reproduction
fragmentation, is thought to be critical in
persistence of populations, especially in ar
impacted by trawling (Le Goff-Vitry and Ro(
2005; Waller and Tyler 2005).

I.A. .a.ii. Solenosmilia variabilis
Duncan, 1873

Description: Solenosmilia variabilis (Fi(
1.4) is a branching coral that often occurs c
secondary constituent of deep-water reefs. It
prominent reef-building species on South Pa


Figure 1.5 The deep coral Enallopsammia profunda. Photo credit: Brooke et al.,

Seamounts, along the Heezen Fracture Zone in
the South Pacific, on Little Bahama Bank, and
south of Iceland (Cairns 1979; Freiwald et al.
2004). S. variabilis is also associated with L.
pertusa, Madrepora spp., and E. profunda in the
western Atlantic on the Blake Plateau and along
the Florida-Hatteras slope (Chapters 7 and 8).

Depth Range: S. variabilis is found at depths
of 220-2,165 m, but is only known to occur at
depths shallower than 1,383 m in the western
Atlantic (Cairns 1979).

Morphology: S. variabilis forms bushy, tightly
branched colonies.

Growth and Age: Limited information is

Reproduction: S. variabilis is a gonochoristic
species with relatively small polyps (3.3 mm),
small oocytes (148 pm), and low polyp fecundity
(290) that increases with polyp size. The species
is thought to be a broadcast spawner with annual
reproduction in late April or May in New Zealand
(Burgess and Babcock 2005).


I.A. 1.b.i. Enallopsammia profunda
(Pourtales, 1867)

Description: Enallopsammia profunda is a major
structure-forming species (Cairns 1979; Rogers
1999). It is often associated with L. pertusa, M.
oculata, and S. variabilis (Reed 2002a; Reed et
al. 2006).

Distribution: E. profunda is endemic to the
western Atlantic and can be found from the
Caribbean to Massachusetts. E. profunda can
contribute significantly to the structure of deep-
water coral banks found at depths of 600-800 m
in the Straits of Florida (Cairns and Stanley 1982;
Reed 2002a). For example, a site on the outer
eastern edge of the Blake Plateau at depths
of 640-869 m contains over 200 coral mounds
where E. profunda is the dominant scleractinian
coral (Stetson et al. 1962; Uchupi 1968; Reed
2002a). Enallopsammia-Lophelia reefs have a
reported maximum vertical relief of 146 m (Reed
2002a; Reed et al. 2006).

Depth Range: E. profunda occurs at depths
from 403-1,748 m (Cairns 1979).

Morphology: This species forms large branching


Figure 1.6 Specimen of Enallopsammia rostrata
(31.4 cm) collected at 1,097 m off Bermuda. Specimen
includes L. pertusa and D. dianthus. Photo credit:
S. Lutz.

colonies up to 1 m in diameter (Cairns 1979;
Freiwald et al. 2004) (Figure 1.5).

Growth, Age, and Reproduction: Limited
information is available.

I.A. 1.b.ii. Enallopsammia rostrata
(Pourtales, 1878)

Description: Enallopsammia rostrata (Figure
1.6) is a widespread scleractinian species that is
known to contribute to the structure of deep coral
reefs. It is reported to form bioherms along the
edges of oceanic banks, such as the Chatham
Rise off New Zealand (Probert et al. 1997). It
is considered a major structure-forming coral in
Hawaii (Chapter 4) and the Caribbean (Chapter

Distribution: E. rostrata has been reported from
eastern and western Atlantic, the Indian Ocean,
and numerous locations in the central and
western Pacific (Cairns et al. 1999), ranging in
latitude from 530 N (in the Atlantic) to 51 S in the
Pacific. In U.S. waters it is the most important
deep-water scleractinian in Hawaii, where it
is common primarily at depths of 500-600 m
(Chapter 4). In U.S. waters of the Atlantic, it
has been reported to occur off Georgia (Chapter
6), Navassa Island and the U.S. Virgin Islands
(Chapter 8).

Depth Range: E. rostrata occurs at depths from
215-2,165 m (Cairns 1979, 1984). In Hawaii, it is

Figure 1.7 Madrepora carolina specimen (27.6 cm)
collected at 333-375 m in the northwest Providence
Channel off Grand Bahama Island. Photo credit:
S. Lutz.

most common at depths of 500-600 m (Chapter
4). It occurs from 300-1,646 m in the western
Atlantic (Chapter 8; Cairns 1979).

Morphology: E. rostrata forms tightly-branched,
bushy colonies (Cairns 1979).

Growth and Age: Adkins et al (2004) reported
that a single colony of E. rostrata from the
North Atlantic was over 100 years old, with an
estimated linear growth rate of 5 mm per year.

Reproduction: Burgess and Babcock (2005)
reported that E. rostrata appeared to be a
gonochoristic, broadcast spawner, although
brooded larvae could not be ruled out. Maximum
oocytes diameter was 400 pm with an average
of 144 oocytes per polyp.


I.A.1 .c.i. Madrepora carolina
(Pourtales, 1871)

Description: Madrepora carolina has been
reported on deep-water reefs, often in association
with E. profunda, and other species, but it is not
known to form the structural framework of these
reefs (Freiwald et al. 2004).

Distribution: M. carolina occurs throughout the
tropical western Atlantic in the Gulf of Mexico
and off the southeastern United States, often co-
existing with M. oculata.

Depth Range: M. carolina occurs from 53-1,003
m, but is most common between 200-300 m
(Chapter 7; Cairns 1979; Dawson 2002).


Figure 1.8a Madrepora oculata coral in
situ, one of the three dominant corals that
make up the deepwater reefs off Florida.
Photo credit: Brooke et al, NOAA-OE,

Morphology: This species forms bush-like
colonies with a thick base up to 28 mm in
diameter (Cairns 1979; Figure 1.7).

Growth, Age, and Reproduction: Limited
information is available.

I.A.1.c.ii. Madrepora oculata
Linnaeus, 1758

Description: Madrepora oculata is not known
to build reefs, but it is typically a secondary
framework builder that occurs among colonies
of L. pertusa off New Zealand, the Aegean Sea,
and northeast Atlantic (Frederiksen et al. 1992;
Freiwald et al. 2004; Waller and Tyler 2005),
among L. pertusa, E. profunda, and S. variabilis
off the southeast Atlantic (Reed 2002a; Reed
et al. 2006) and G. dumosa off New Zealand
(Cairns 1995). Recent molecular studies of the
scleractinians have given a new insight into the
evolutionary history of this group. Analysis of
mitochondrial 16S rDNA suggests that M. oculata
may have been misclassified, and it may actually
form a monotypic clade between the families
Pocilloporidae and Caryophylliidae (Le Goff-Vitry
et al. 2004).

Distribution: M. oculata is one of the most
widespread deep-water coral taxa. It has been
recorded in temperate and tropical oceans
around the world, extending from 690 N off

Figure 1.8b Madrepora oculata sample
collected at the Lophelia coral banks
off the coast of South Carolina. Photo
Credit: Ross et al. and NOAA-OE.

Norway to 590 S latitude in the Drake Passage.
Large individual colonies of M. oculata occur on
exposed hard substrate throughout the Gulf of

Depth Range: This species is known to occur
from 55-1,950 m (Zibrowius 1980; Cairns 1982).

Morphology: M. oculata has a complicated
skeletal morphology. It has extremely variable
morphology, forming large bushy or flabellate
colonies with a massive base that can be
several centimeters in diameter (Cairns 1979).
Colony branches have very distinctive "zig-zag"
morphology (sympodial branching; Figures 1.8a
and 1.8b). M. oculata is reported to be more
fragile than L. pertusa, limiting its structure-
building capability.

Growth and Age: Limited information is

Reproduction: The reproductive ecology of M.
oculata contrasts sharply with that observed in
Lophelia. While both are gonochoristic broadcast
spawning species, M. oculata is thought to
produce two cohorts per year and the oocytes
are more than 2.5 times larger than L. pertusa


Figure 1.9 Oculina varicosa in the Oculina HAPC. Phot(
L. Horn, NOAA Undersea Research Center at UNC-

(max = 405 mm diameter), but the fecundity is
much lower (a total fecundity of 10- 60 oocytes
per polyp vs. 3,000 oocytes for L. pertusa; Waller
and Tyler 2005).

I.A.1.c.iii. Oculina varicosa
Lesueur, 1821

Description: Oculina varicosa (the ivory tree
coral) is an important deep reef-building species
that forms thickets of large branched colonies
along the eastern Florida shelf.

Distribution: 0. varicosa is restricted to the
western Atlantic, including the Caribbean and
Gulf of Mexico, Florida to North Carolina and
Bermuda (Verrill 1902; Reed 1980). The deep-
water Oculina reefs, however, are only known off
the east coast of central Florida at depths of 70-
100 m (Avent et al. 1977; Reed 1980, 2002b),
occurring as offshore banks and pinnacles up to
35 m in height (Reed 2002b; Reed et al. 2005)
(Figure 1.9).

Depth Range: Depth range of 0. varicosa has
been reported from 2-152 m (Verrill 1902; Reed
1980). It is an unusual coral in that it occurs
in both shallow and deep waters (Reed 1981),
and is facultatively zooxanthellate, containing

symbiotic algae only in shallow waters
(2-45 m).

SMorphology: There are morphological
differences between the shallow and
deep-water colonies of 0. varicosa.
,* Shallow populations (2-45 m) are
dominated by stout, thickly branched
colonies, possibly in response to
wave action (Verrill 1902; Reed 1980).
Deeper colonies (49-152 m) are more
fragile and taller than their shallow
counterparts, with colonies growing up
to 2 m in diameter and height (Reed
1980, 2002b).

Growth and Age: The linear branch
growth rate of 0. varicosa appears to
be faster in deeper water (16 mm yr1 at
80 m) where zooxanthellae are absent,
than at 6 m depth (11 mm yr1). These
credit: differences may be due to environmental
factors such as greater sedimentation
rates and more variable temperature
extremes, as well as morphological
differences in which shallow colonies
put more energy into diameter than height (Reed
1981, 2002b).

Reproduction: 0. varicosa is a gonochoristic
broadcast spawning species, producing large
numbers of small eggs which are released
annually in August and September (Brooke and
Young 2003). Planulae have a relatively long

Figure 1.10 Madracis myriaster specimen (30.2 cm)
collected from 200 m off Jamaica. Photo credit:
S. Lutz


U.S. waters it occurs in the Gulf
of Mexico, Straits of Florida,
off the Atlantic coast of Florida
and Georgia, and in the U.S.
Caribbean off Puerto Rico and
the U.S. Virgin Islands.

Depth Range: M. myriaster is
found at depths ranging from
37-1,220 m (Chapter 8; Cairns

Morphology: M. myriaster is
a branching species that forms
broad, bushy colonies of 30-40
cm in height (Cairns 1979).

Growth, Age,
information is available.


I-igure 1.11 (ola coral (ceraraia sp.) in Hawall wtn a pu
Clavularia grandiflora growing on it. Photo credit: A. Bac

planktonic period (at least 22 days) (Brooke
and Young 2003, 2005), which provides the
potential for widespread transport between
deep reef tracks as well as cross-shelf transport
(Smith 1983). This strategy may help facilitate
recovery of degraded areas, although very little
coral recruitment has been observed to date in
damaged areas (Brooke and Young 2003).


I.A.1.d.i. Madracis myriaster
(Milne-Edwards and Haime, 1849)

Description: Madracis myriaster (Figure 1.10)
is a deep-water species in the predominantly
shallow-water family Pocilloporidae. It is
reported as a primary framework-builder of
Caribbean deep coral banks off Colombia (Reyes
et al. 2005). It is considered a major structure-
forming coral in the southeast U.S. (Chapter 6)
and the Caribbean (Chapter 8).

Distribution: M. myriaster is endemic to the
tropical northwestern Atlantic Ocean (Cairns
et al. 1999), between 70 and 290 N latitude. In


Zoanthids are colonial, sea
anemone-like anthozoans,
rple octocoral mostly occurring in shallow
tropical waters. While most of
the more than 100 species of
zoanthids do not form skeletal structures, deep-
water gold corals are one taxon found in this
order that does form rigid skeletons and grows
to large sizes.


I.A.2.a.i. Gerardia spp. (Gold corals)

Description: Gerardia spp. form branching
colonies that have an axis of dense, hard
proteinaceous material. The skeleton of gold
corals is used in the manufacture of coral
jewelry. Gold corals were harvested from the
Makapu'u Bed off Hawaii between 1974 and
1978 (Chapter 4). The taxonomy of this group is
not well defined.

Distribution: Gold corals in the family
Gerardiidae are found on hard substrates such
as basalt and carbonate hardgrounds. These
forms of substrate are common on seamounts
in the north and equatorial Pacific and Atlantic

Depth Range: In U.S. waters, gold corals have





idLt dcllieve relatively larye sizes bvvaiuliy (klviev1 ku r\t./-\LO)
id Auster 2005). Soft corals of the genus
unephthea (formerly Gersemia) are widespread Description: Isididae is a large family
id are the most abundant corals in the Bering 150 species of mostly deep-water co
ea (Chapter 2). True soft corals of the order most common deep-water genera ar
Icyonacea generally lack a rigid internal skeleton Isidella and Keratoisis. Acanella ai

Stoloniferans, now included in the Alcyona
have small polyps that are often connect
each other by a thin runner or stolon. With
exception of the tropical shallow-water or!
pipe coral (Tubipora musica, most are
important structure-forming corals. How(
a few species can form extensive mats
hard surfaces such as rocks, other corals,
sponges (Stone 2006).


Major structure-forming families in the c
Gorgonacea include Isididae, Corallii
Paragorgiidae, and Primnoidae (Morgan e
2006), with species in the families Plexauri
Acanthogorgiidae, Ellisellidae, Chrysogorgii
and Anthothelidae providing structure to s
degree (Appendix 1.1 and 1.2). At leas
families are known to occur in waters de,
than 200 m (Etnoyer et al. 2006). Gorgon
are the most important structure-forming cc
in the Gulf of Alaska and the Aleutian Isla
where they form both single- and multi-spe
assemblages (Chapter 2). For exarr
Primnoa pacifica forms dense thickets in
Gulf of Alaska (Krieger and Wing 2002), while
many as 10 species are found in Aleutian Is
coral gardens (Stone 2006). Most gorgon
have a solid proteinaceous (gorgonin) ce
axis with embedded calcareous sclerites
provide support, covered by a thin layer of ti.
(coenenchyme and polyps) with embec
calcareous spicules (Fabricius and Alders
2001). They often exhibit a branc
morphology, can occur at high density and cc
and reach considerable size (>3 m tall),
providing structure and habitat for associ

a, in mud rather than on hard substrata (Morten
to and Buhl-Mortensen 2005a). Several spe
ie are collected for jewelry.
)t Distribution: Bamboo corals are though
'r, have a cosmopolitan distribution and impor
in structure-forming species have been ident
id in the Gulf of Mexico, the Southeast, Ha\
the West Coast, the northeast Pacific and Ir
Pacific (Fabricius and Alderslade 2001; Etn(
and Morgan 2003; Appendix 1.1 and 1.2).

Depth Range: In general bamboo corals oi
er below 800 m (Etnoyer and Morgan 2005),
B, the deepest recorded at 4,851 m (Bayer
II. Stefani 1987). In Alaska bamboo corals
B, observed between 400 and 2,827 m but h
B, been collected from depths of 3,532 m (Cha
ie 2). However, four genera have been repo
2 from tropical Indo-Pacific reefs at depths of
er 120 m (Fabricius and Alderslade 2001).
Is Morphology: Colonies can be whip-like but
s, usually branched, bushy or fan-like (Figure 1
is and can range in size from tens of centime
B, to over a meter (Verrill 1883). Colonies ha\
ie distinctly articulated skeleton of heavily calc
is internodes and proteinaceous gorgonin nc
id (a stiff leathery matrix consisting of protein
is mucopolysaccharides). The alternating segmi
al give the isidid branches a unique bamboo
at appearance (Figure 1.13), hence the n,
ie "bamboo-coral."
le Growth and Age: Recent studies by Andr
ig et al. (2005a,b) have estimated radial grc
'r, rates for bamboo corals that ranged f
is approximately 0.05 (age 150 years) to 0.117
'd yr-1 (age 43 years). Linear growth rates up t(
mm yr1 have been estimated for Lepidisis sl
New Zealand waters (Tracey et al. in press).

Reproduction: Reproductive strategy is thot
to be similar to that of other octocorals
colonies having separate sexes and gami


Figure 1.13 A. Bamboo coral (Keratoisis
sp.) on the Davidson Seamount (1,455
meters). Coral colony age estimates ex-
ceed 200 years. Image courtesy of NOAA,
MBARI 2002. B. Close-up view of white
bamboo coral (Keratoisis flexibilis) show-
ing the coral's extended feeding polyps.
This coral and other filter feeders orient so
that they are perpendicular to the current,
positioning themselves to be in the flow of
food carried in the current. Photo credit:
Brooke et al., NOAA-OE, HBOI.

being broadcast into the water column in a
synchronous manner (Fabricus andAlderslade


Description: The family Coralliidae was recently
divided into two genera Paracorallium and
Corallium (Bayer and Cairns 2003). The only
known populations of pink and red corals large
enough to support commercial harvest are found
north of 190 N latitude, including seven species
harvested in the western Pacific and one

collected in the Mediterranean. All species of
Corallium identified in the Southern Hemisphere
occur in low abundance (Grigg 1993). The
family Coralliidae contains the most valuable
taxa of precious corals. It is traded in large
quantities as jewelry and other products, and as
raw coral skeletons. Of the 31 known species
in this family, seven are currently used in the
manufacture of jewelry and art (Cairns in press;
Figure 1.14a). One species, Corallium rubrum,
has been harvested for at least 5,000 years from
the Mediterranean. Other species have been
harvested for 200 years in the western Pacific
off islands surrounding Japan, Taiwan, and the


Vk~h~~ 3~""~

Philippines, and for 40 years in the western
Pacific off Hawaii and international waters around
Midway Islands (Grigg 1993).

Distribution: The family is widely distributed
throughout tropical, subtropical, and temperate
oceans including five species from the Atlantic
Ocean, one from the Mediterranean Sea, two
from the Indian Ocean, three from the eastern
Pacific Ocean, and 15 from the western Pacific
Ocean (Grigg 1974; Weinberg 1976; Cairns in
press). In U.S. waters, they are best known from
banks off Hawaii (Chapter 4). They have also
been found on seamounts in the Gulf of Alaska
(Baco and Shank 2005; Heifetz et al. 2005),
Davidson Seamount off the California coast
(DeVogeleare et al. 2005), and the New England
Seamounts in the Atlantic (Morgan et al. 2006;
Figure 1.14b).

Depth Range: Depths for this family range from
7 m to 2,400 m (Bayer 1956; Weinberg 1976).

Morphology: Corallium spp. have a hard
calcareous skeleton with an intense red or
pink color (Figure 1.14a). They are sedentary
colonial cnidarians with an arborescent growth
form, attaining heights ranging from 50-60 cm
(C. rubrum) to over 1 m (U.S. Pacific species).

Growth and Age: Corallium species are very
slow growing, but individual colonies can live
for 75-100 years. For example, C. rubrum
exhibits average annual growth rates of 2-20
mm in length and 0.24-1.32 mm in diameter.
Corallium secundum, a commercially valuable
species found off Hawaii (Chapter 4), is reported
to increase in length at rates of about 9 mm
yr1 (Grigg 1976). Natural mortality rates of C.
secundum vary between 4-7%, with turnover
of populations occurring every 15 to 25 years
(Grigg 1976).

Reproduction: Aspects of reproductive
biology have been studied for C. rubrum and C.




secundum only. These species have separate
sexes and an annual reproductive cycle. C.
rubrum reaches maturity at 2-3 cm height and
7-10 years of age2 (Santangelo et al. 2003;
Torrents et al. 2005); C. secundum reaches
maturity at 12 years (Grigg 1993). Usually, C.
rubrum is a brooder with a short-lived passive
larval stage while C. secundum is a broadcast
spawner. Planulation occurs once per year,
primarily during summer. Larvae remain in the
water column for a few days (4-12 days in the
laboratory) before settling in close proximity to
parent colonies (Santangelo et al. 2003).

21n earlier studies, more than 50% of colonies were re-
ported to reach sexual maturity at 2 years, and all colonies
over 5 years were fertile. Recent aging studies suggest
that these reports underestimated the true age of reproduc-
tive maturity by 3-4 years (Marschal et al. 2004).

re 1.15 a. Paragorgia, or "bubblegum corals," grow to
2.5 m tall. On Davidson Seamount, where this photo
aken, they are found primarily on the highest
tions. b. Close up of Paragorgia. Photo credit: The
Ison Seamount Expedition, MBARI, and NOAA-OE.

Description: The small family Paragorgiidae,
commonly referred to as "bubblegum corals," has
recently been expanded to includes nine known
species in the genus Paragorgia, (Sanchez
2005). These corals are large branching
gorgonians (Figure 1.15) and are thought to
reach the largest size of any sedentary colonial
animal. For example, colonies of Paragorgia
arborea in New Zealand have been reported to
reach 10 m in height (Smith 2001).

Distribution: P. arborea has been reported to
have a bipolar distribution, occurring in deep
waters of the Southern Hemisphere and in the
North Atlantic and Pacific Oceans (Grasshoff
1979). In the U.S., P arborea, occurs in the
submarine canyons off Georges Bank at depths


Figure 1.16. Large primnoid coral with associated brittle stars on Dickinson Seamount, Gulf of Alaska. Photo
credit: The Gulf of Alaska Seamount Expedition, and NOAA-OE.

thickets. It is also reported to be common in the
Aleutian Islands of Alaska (Chapter 2; Etnoyer
and Morgan 2003) and on Alaskan seamounts
and Davidson Seamount off California. Recent
analysis of specimens of Paragorgia arborea
collected off the Atlantic coast of Canada and a
morphologically similar Paragorgia sp. from the
Pacific were genetically dissimilar (Strychar et al.

Depth Range: Paragorgiids have been found
in the Pacific at depths ranging from 19-1,925
m (Etnoyer and Morgan 2003). In the northeast
Atlantic they have been found to depths of 1,097
m (Mortensen and Buhl-Mortensen 2005a).

Morphology: Like other gorgonians, colonies
have a proteinaceous skeleton with embedded
spicules covered in a soft tissue, and polyps
have eight feather-like tentacles. P arborea
exhibits distinct intraspecific color variation, with
red, pink, orange, and white morphs reported.

Growth and Age: Growth rates of bubblegum
coral are not well defined. Mortensen and Buhl-
Mortensen (2005b) report estimates of linear
growth rates for P arborea in New Zealand
and Norway between 2.2-4.0 cm yr1 and 0.8-
1.3 cm yr1 respectively. Andrews et al. (2005a)
estimated a Pacific species to have grown
at a minimum of 0.5 cm yr1 based on a single
observation of a 20 cm coral on a telegraph
cable submerged for 44 years.

Reproduction: Reproductive strategy is thought
to be similar to that of other octocorals with
colonies having separate sexes and gametes
being broadcast into the water column in a
synchronous manner (Fabricus and Alderslade


Description: Primnoidae is a large family (>200
species) that includes a number of conspicuous
and abundant branching species in the genera


Primnoa (the red tree corals) and Callogorgia.
These corals attach to rocky outcrops and
boulders in the presence of strong currents.

Distribution: They are among the most common
large gorgonians, occurring in dense thickets in
some regions and, in the U.S., appear to reach
their highest abundance in Alaska (Figure 1.16;
Chapter 2; Etnoyer and Morgan 2005).

Depth Range: Etnoyer and Morgan (2003) found
the northeast Pacific depth range for Primnoidae
to be 25-2,600 m, with the majority occurring
shallower than 400 m. Primnoa resedaeformis
occurs from 91-548 m in the northwestern
Atlantic, where it is among the most abundant
species (Cairns and Bayer 2005).

Morphology: Primnoa spp. form a branching
tree-like structure with a skeleton composed of
calcite and a hornlike protein called gorgonin.

Growth and Age: Primnoa spp. can reach over 7
m in height (Krieger 2001). Growth of deepwater
primnoids is slow, with growth rates estimated at
1.60-2.32 cm in height and 0.36 mm in diameter
per year for a Primnoa sp., found in the Gulf of
Alaska (Andrews et al. 2002)3. Mortensen and
Buhl-Mortensen (2005b) estimated growth rates
for P resedaeformis in the Canadian Atlantic,
at 1.8-2.2 cm per year for young colonies (<30
years) and 0.3-0.7 cm per year for older colonies,
with a maximum age of 60 years fro the sampled
corals. Risk et al. (1998, 2002) estimated large
colonies of P resedaeformis, sampled from Nova
Scotia to be hundreds of years old.

Reproduction: Reproductive strategy is thought
to be similar to that of other octocorals with
colonies having separate sexes and gametes
being broadcast into the water column in a
synchronous manner (Fabricus and Alderslade


Description: Pennatulaceans are a diverse but
poorly known group of octocorals that include
16 families, most of which live in the deep sea.

3The authors identified this species in Alaska as Primnoa
resedaeformis, which is now thought to occur only in the

They are uniquely adapted to soft-sediment
areas (Figure 1.17); many are able to uncover
themselves when buried by shifting sands and
re-anchor when dislodged. They burrow and
anchor by means of peristaltic contractions
against the hydrostatic pressure of the peduncle
(Williams 1995). Certain species, such as
Ptilosarcus gurneyi, are capable of completely
withdrawing into the sediment (Birkeland
1974). Though pennatulaceans are common
in many parts of the U.S. Exclusive Economic
Zone (EEZ) their contribution as habitat and
to diversity of associated species is not well
documented or understood. Brodeur (2001)
found dense aggregations of rockfish (Sebastes
alutus) associated with "forests" of the sea pen
Halipteris willemoesi in the Bering Sea. They
are probably not structure-forming in the same
sense as other coral groups discussed above
but they may provide important habitat in areas
that cannot be colonized by structure-forming
gorgonian and scleractinian corals. Given their
widespread distribution in soft-sediment areas,
sea pens may be the most abundant deep coral

Distribution: Sea pens are found in all the
world's oceans (Fabricius and Alderslade 2001).
In Alaskan waters a few species are known to
form extensive groves (Figure 1.17). The South
Atlantic Bight and Gulf of Mexico appear to be
relatively depauperate in comparison to other
U.S. regions (Chapters 6 and 7).

Depth Range: Pennatulaceans are known
from shallow waters to the abyssal plains. Most

Figure 1.17 Dense groves of the sea pen Ptilosarcus
gurneyi are found on soft-sediment shelf habitats
in the Gulf of Alaska and Aleutian Islands. Photo
credit: P. Malecha, NOAA's National Marine Fisheries


pennatulaceans live in the deep
sea, and the deepest known
corals are sea pens in the genus
Umbellula, which have been
recorded at depths greater than
6000 m (Williams 1999).

Morphology: The sea pens
and sea pansies (order
Pennatulacea) differ from other
octocorals in that there is a
large, primary polyp that gives
rise to secondary dimorphic
polyps (autozooids and
siphonozooids), and a stem-
like base or foot pedunclee)
that is anchored in the sand.
Many species are elongate and
whip-like and are supported
by an internal calcareous axis.
Umbellula, a genus found on
the Atlantic abyssal plain and
deep-water areas of the North
Pacific, is characterized by a
long stalk that can be a meter
or more in length, with a series
of secondary polyps mounted at
the end; another species from
Alaska, H. willemoesi, attains
a height greater than 3 m (R.
Stone, personal observation).

Growth and Age: Wilson et al.
(2002) report growth rates for
H. willemoesi of 3.6 to 6.1 cm
yr1 depending on size with an
estimated longevity approaching Figure 1.18 Stylaster coral. Photo credit:
50 years for moderately sized Oceanlight, Carlsbad, CA.

Reproduction: Sea
broadcast spawners
information is limited.

pens are known to be
but other reproductive


Hydrozoans are a mostly marine group
of cnidarians including the hydroids and
hydromedusae (e.g., jellyfish). Most hydrozoans
alternate between a polyp and a medusa stage.
One order, Anthoathecatae, contains species
with calcium carbonate skeletons and are
classified as coral-like organisms: the stylasterid

corals (suborder Filifera, family Stylasteridae),
which include many deep-water species, and
the shallow-water fire corals (suborder Capitata,
family Milleporidae).


I I.A.1.a Family Stylasteridae
(Stylasterid corals)

Description: The order Anthoathecatae,
previously identified as Filifera (now suborder
Filifera) or Stylasterina, contains the stylasterid
corals (also known as lace corals) in the family


Stylasteridae, a group of calcareous encrusting
or branching colonial species. Stylasterids
are often confused with stony corals due to
their calcareous nature but the resemblance is

Distribution: As a group, the stylasterid corals
occur worldwide and at a wide range of depths.
About 90% of the 250 extant stylasterid species
live exclusively in deep waters occurring as deep
as 2,700 m (Cairns 1992a). Other species,
such as the Pacific Stylaster sp. (Figure 1.18),
may occur in relatively shallow water. Stylaster,
Distichopora, and Pliobothrus are among the
better-known genera in this family. In U.S. waters
they have been reported from most regions
except the northern Gulf of Mexico (Cairns
1992b), but appear to be of particular importance
as structure forming components in the Straits of
Florida (Chapter 7) and Alaska (Chapter 2).

Depth Range: Stylasterids may be found in
shallow-water reef systems in only a few meters
of water; deep-water species occur from 79-
2,700 m depth (Broch 1914; Cairns 1992a;
Etnoyer and Morgan 2003).

Morphology: Stylasterids have a hard calcium
carbonate skeleton covered with thousands
of pinhole-sized pores. Each pore contains a
single gastrozooid, a stout feeding polyp with
eight tentacles. These are surrounded by 5-9
dactylozooids, which are thin sensory/stinging
polyps devoid of tentacles but armed with
batteries of nematocysts. Though they have
a wide range of morphological forms, most
have extremely fragile branches and are highly
susceptible to physical damage. In Alaska some
species, e.g.,Stylaster cancellatus, may grow
to almost one meter in height (R. Stone, pers.

Growth and Age:

Limited information is

Reproduction: Stylasterids are usually
gonochoristic and fertilization is internal. The
male and female reproductive structures or
gonophores, develop in epidermally lined cavities
called ampullae (rounded inclusions in the
calcareous skeleton). These ampullae usually
appear as small hemispheres on the surface
of the colony, but occasionally are completely
submerged in the calcareous coenosteum. The

larvae develop in the gonophores and leave via
small pores near the ampullae (Ostarello 1973,
1976). Larvae of the species studied to date are
short-lived and non-dispersive (Ostarello 1973),
which has implications for ecosystem recovery
from disturbance (Brooke and Stone in press).


Structure-forming deep corals are generally
slow growing and fragile, making them and
their associated communities vulnerable to
human-induced impacts, particularly physical
disturbance. With the exception of a few areas
(e.g., the Oculina Banks), the extent of habitat
degradation resulting from these threats is
largely unknown although there is increasing
information on significant impacts in some areas.
Activities that can directly impact deep coral
communities include fishing using bottom-tending
fishing gear, deep coral harvesting, oil and gas
and mineral exploration and production, and
submarine cable/pipeline deployment. Invasive
species, climate change and ocean acidification
represent additional serious threats. Though
not exhaustive, this list does include the more
important activities that may alter deep coral
habitat. The extent of impact from these activities
and the type of stressors that cause the most
degradation vary among regions. For example,
impacts from mobile bottom-tending fishing gear
are the largest potential threat in many areas of
Alaska, but are not an issue in the U.S. Pacific
Island regions where trawling is banned. In
Hawaii, the harvest of certain deep coral species
- including black corals, pink coral, gold coral,
and bamboo corals is permitted and regulated
by the Precious Coral Fishery Management
Plan (FMP) in federal waters and under Hawaii
Administrative Rules in state waters.

Bottom Trawling and Other
Bottom-tending Fishing Gear Impacts

A number of different types of fishing gear impact
the seafloor and pose potential threats to deep
coral communities. Table 1.2 lists different types
of fishing gear known to impact deep corals, a
description of the gear, their impacts, and the
level of severity as a result of these interactions.
Bottom trawling is the largest potential threat
to deep coral habitat for several reasons: the


Table 1.2 Major bottom-contact fishing gear types used in U.S. fisheries and a description of their potential
impact on structure-forming deep corals. This table identifies only the potential severity of disturbance to
deep corals if they are encountered by the gear based on reported instances of interactions. It is not meant
to indicate that interactions between these gears and deep corals currently occur in the U.S. EEZ. Regional
Fishery Management Councils have analyzed potential gear impacts and have proposed measures to
minimize to the extent practicable adverse impacts of these gears on essential fish habitat (see Section on
U.S. Conservation and Management Measures). **NRC 2002; t High 1998; NMFS 2004; t Eno et al. 2001;
, Stone et al. 2005.

Major types
of bottom-
fishing gear

Gear types that
may impact
deep corals

Description of gear

Description of impact on

A large net is held open by two doors
and dragged behind a fishing boat Incidentally removes, dis-
Incidentally removes, dis-
Bottom/ along the seafloor; gear in contact laces ordama
otter trawls with seafloor can be 30-100 m in o*
co rals**
length; primarily used to harvest
Trawls demersal finfish and rock shrimp
Similar type of gear to the bottom May come in contact with
trawl but designed to harvest pelagic bottom during fishing,
Mid-water trawls
fish species; no protective gear on which can remove, dis-
the footrope place or damage corals
A large steel frame is dragged behind Incidentally removes,
Dredges Scallop dredges a fishing boat along the seafloor; displaces or damages
specifically used to harvest scallops corals I


Traps or

Bottom-set long-
demersal long-

Single-set pots

Longline pots

A nylon or poly line with up to
thousands of attached hooks; de-
ployed along the seafloor in lengths
up to 2-5 km
A pot or trap that is constructed of
wooden slats or coated wire mesh;
set as a single pot on the bottom to
harvest finfish or shellfish; pots vary
in size up to 4.5 m2

Single pots are strung together on a
long line (10-90 pots)

May entangle or detach
corals during retrieval t

Limited spatial damage to
corals during pot retrieval

Damages corals during
pot retrieval and entan-
glement with lines; under
certain conditions gear
can be dragged like a
plough on seafloor

J _____________________________________ -

of distur-




area of seafloor contacted per haul is relatively
large, the forces on the seafloor from the trawl
gear are substantial, and the spatial distribution
of bottom trawling is extensive. Although not as
destructive as bottom trawls and dredges, other
types of fishing gear can also have detrimental
effects on deep coral communities. Bottom-set
gillnets, bottom-set longlines, pots and traps all
contact the benthos to some degree. Vertical
hook and line fishing, used in both recreational
and commercial fishing, has the potential for
some damage to fragile corals by the weights
used, but such damage is likely to be minimal
compared to other bottom-tending gear (NRC

2002; Kelley and Ikehara 2006). Chuenpagdee
et al. (2003) surveyed U.S. fishery management
council members (including fishers), scientists
who served on the National Research Council's
Ocean Studies Board or its study panels, and
fishery specialists of conservation organizations,
on their opinions of the ecological impacts of
various classes of fishing gear. There was
general agreement among respondents that
unmitigated impacts to biological habitat from
dredge and trawl gear were expected to be more
severe than those of other gears.






I ...






mnere nave oeen nimitea stuaies on mne long- riyure I.Lu oieei puis a1e useu LU IIiaves~ nialiy spe-
term impacts of trawling and dredging (FAO cies of crabs in Alaska. Some pots, such as this one,
2005). Recovery rates range from one to five measure 2 x 2 x 1 m and may weigh more than 300 kg.


Figure 1.21 The black coral divers of Lahaina, Maui.
Team leader Robin Lee is the diver wearing the cap.
Photo credit: R. Grigg.

than that of mobile bottom-tending gear, since
the extent of habitat impacted is much more
limited. The potential for significant disturbance
is greater if pots or traps are dragged along the
bottom during retrieval (Freiwald et al. 2004;
Figure 1.20). Gorgonians (Primnoa spp.) were
reported to disappear in an area where prawn
pots were set because of coral entanglement
in the mesh of the pots (Risk et al. 1998). In
certain fisheries, numerous traps are connected
in series. These "longline pots" can cause a high
level of disturbance to deep coral communities
while the other types of bottom-contact gear
cause moderate levels of disturbance (Table 1.2
and Chapter 2).

Demersal longlines and gillnets
High (1998) recognized that bottom contact
by gillnets can alter the seafloor and that large
branching corals can be detached, entangled,
and brought up as bycatch by longlines. In
Nova Scotia fishermen reported the snagging
of gorgonian corals when longline gear became
tangled (Breeze et al. 1997), and Mortensen et
al. (2005) identified direct and indirect impacts
of longlines on corals off Canada. Fishing gear
fixed to the seafloor with anchors and weights,
such as gillnets, also has the potential to
impact fragile deep coral habitat, as reported off
Porcupine Bank in the northeast Atlantic (Grehan
et al. 2004).

Certain deep corals have been collected or
harvested for jewelry and curios since antiquity.
In the U.S., commercial harvest of precious

corals4 for jewelry and curios has occurred
off Hawaii periodically since 1958. In federal
waters, precious corals are managed under
the Precious Corals Fishery Management Plan
(FMP). Implemented in 1983, this FMP regulates
two distinct fisheries: one for black corals and
one for all other precious corals. The Precious
Corals FMP provides regulations on permits,
prohibitions, seasons, quotas, closures, size
restrictions, areas restrictions, including recent
prohibitions on non-selective harvest.

In state waters, black and pink corals are
managed under the Hawaii Administrative Rules.
While black coral has been harvested in both
federal and state waters for nearly five decades,
harvest of precious corals (mostly Corallium
spp.) in U.S. waters only occurred from 1966
to 1969 and from 1972 to 1979, with a short
revival of the fishery in 1999 and 2000. Due to
the low abundance of precious corals and their
vulnerable life history traits, coral harvesting may
be the single largest impact to deep corals in
Hawaii. The sustainability of black coral harvests
is currently of greatest concern, as populations
are also being impacted by a rapidly spreading
invasive coral, Carijoa riisei (see invasive species
discussion below and Chapter 4).

In 1990 the Gulf of Mexico Fishery Management
Council approved an amendment under the Coral
and Coral Reefs FMP allowing harvest of up to
50,000 octocoral colonies (with the exception of
sea fans) per year, for commercial trade in the
aquarium industry. In Alaska, a fishery for corals
was proposed but never developed, even though
this region contains a variety of precious corals
that are harvested in other regions.

Mineral Resource Exploration and Extraction
Oil and Gas Exploration and Production
Exploration for and production of oil and gas
resources can impact deep coral communities
in a variety of ways. Potential threats include
the physical impact of drilling, placement of
structures on the seafloor (e.g., platforms,
anchors, pipelines, or cables), discharges
from rock-cutting during the drilling process,
and intentional or accidental well discharges

4Precious corals refer to red and pink corals (family Coral-
liidae). The term is often used more broadly to also include
black corals (family Antipathidae), gold corals (family Ge-
rardiidae), and bamboo corals (family Isididae).


Acnve leCses oy water aepin
j < 1,000 ft so 0 so
1,000 1,499 ft
S1,500 4,999 ft 50 0 50m A
S5,000 7,499 ft
S>7,500 ft
Figure 1.22 Map of the Gulf of Mexico showing active leases by water depth in 2005. Image credit: Minerals
Management Service.

or release of drilling fluids. Deep-water drilling
requires special synthetic-based fluids that
operate at low water temperatures. These fluids
are not highly toxic, but accidental release could
also include oil and large amounts of sediment.
While oil contains toxic fractions, a large spill
of unconsolidated sediments could smother
nearby corals. Smothering and death of corals
by drilling muds and cuttings was observed on
Lophelia pertusa colonies living on an oil platform
in the North Sea close to drilling discharge points
(Gass and Roberts 2006).

The use of anchors, pipelines, and cables for
oil exploration/extraction can be destructive to
sensitive benthic habitats as well. Evidence
of damage from a wire anchor cable during
oil and gas drilling was seen during a survey
of the northeastern Gulf of Mexico, including
severe damage to corals within a 1-1.5 m swath
(Schroeder 2002). Cables associated with oil
drilling activities have the largest potential impact
on deep coral communities in the Gulf of Mexico
and in some areas of the West Coast and Alaska
regions. There is increasing interest in laying
liquid natural gas pipelines across the East
Florida shelf, and there is a potential for damage
to the deep coral communities in the area.

Oil and gas exploration and production currently
occur in the Gulf of Mexico, Alaska, and the West
Coast regions. The spatial scale of exploration
varies among these regions. Approximately
98% of all active leases occur in the Gulf of
Mexico (8,140 leases; Figure 1.22) and the other
2% occur off southern California (79 leases),
northern Alaska in the Arctic Ocean (-64 leases),
and southern Alaska in Cook Inlet (~2 leases).
In 1995, the Deep Water Royalty Relief Act
allowed oil exploration and production to move
into deeper waters of the Gulf of Mexico and
leasing activity increased exponentially for this
region. Since 2001 deep-water leasing activity
has leveled off but there are still over 4,000
active leases operating in the Gulf of Mexico at
depths greater than 300 m (Figure 1.22; French
et al. 2005). The movement of leasing activity to
deeper waters could have a significant impact on
important structure-forming deep corals such as
L. pertusa. Leases are active for five years and,
while they do not necessarily lead to oil and gas
extraction, they often include activities such as
exploratory drilling and other activities that may
be detrimental to corals.

Sand and Gravel Mining
Sand and gravel mining usually occurs within
state waters in relatively shallow areas, but


interactions with deep coral habitat from this
industry may be more common in the future.
In the past 10 years, to offset extensive beach
erosion along the U.S. East Coast, 23 million
cubic yards of sand was mined from the Outer
Continental Shelf (OCS). While most OCS sand
mining projects to date have occurred on ridges
and shoals off the costs of the eastern U.S. and
Gulf of Mexico (MMS 2003), 14 states (Alabama,
California, Delaware, Florida, Louisiana, Maine,
Maryland, Massachusetts, New Hampshire, New
Jersey, North Carolina, South Carolina, Texas,
and Virginia) are working cooperatively with the
Minerals Management Service (MMS) to identify
sand mining sites within the OCS to replenish
coastal areas in the future5. Sand mining
activities tend to be highly localized and primarily
impact soft-bottom communities.

OCS sand-mining areas do not tend to overlap
with deep coral habitat because most deep
structure-forming corals are found on hard-
substrates. However, sand mining activities
could impact sea pen groves or indirectly affect
other nearby deep coral communities through
an increase in sedimentation. The regions that
would most likely be impacted by sand mining
are the Northeast, Southeast, Gulf of Mexico,
and the Central West Coast.

Seafloor Mining
Mining the deep seafloor for metals is not yet
a commercially viable enterprise. Historically,
interest has focused on the prospect of mining
manganese nodules that are formed at abyssal
depths. But other potential resources are
being considered. Interests include cobalt-
enriched crusts, which occur in a thin layer on
the flanks of volcanic islands and seamounts
at depths of 1,000 to 2,500 m locations and
depths that also include deep corals. Massive
sulfide deposits on the seafloor appear to be an
even more promising mining resource. These
deposits contain copper, gold, zinc, and silver
associated with extinct hydrothermal vents,
and may yield up to 40 times the resources of
land-based mines (Schrope 2007). Important
deep coral communities have not been reported
in association with seafloor massive sulfide
deposits (ISA 2007).

'More information on this program is available online at

Lonventon aaminisier mineral resources on
and below the seabed in areas beyond national
jurisdiction. Potential environmental impacts
associated with mining cobalt-enriched crust and
seafloor massive sulfide deposits were recently
reviewed in an International Seabed Authority
workshop (ISA 2007).

Submarine Cable/Pipeline Deployment
Deployment of gas pipelines and fiber optic
cables can cause localized physical damage to
deep corals. MMS regulations state that gas
pipelines placed at depths greater than 61 m
are not required to be anchored to the bottom or
buried in the substrate. In these deeper habitats
the pipelines are placed by using 12 anchors
connected to a barge that are "walked" across
the seabed, potentially causing damage through
direct physical contact and via the swath of
the anchor chains (detailed in Koenig et al.
2000). Pipelines and associated structures not
anchored to the bottom may be moved around
by currents and continue to damage nearby coral
communities. For example, a wire anchor cable
used during oil and gas drilling caused severe
damage to corals within a 1-1.5 m swath area
in the northeastern Gulf of Mexico (Schroeder
2002). Cable impacts associated with oil drilling
activities would be most common in the Gulf of
Mexico as well as some areas of the West Coast
and Alaska regions. Fiber optic cables are a
minimal threat to deep coral communities off the
West Coast. Most of these cables are buried in
the sediment, with localized impacts to deep sea
communities occurring during installation, e.g.,
increased sedimentation, etc. (Brancato and
Bowlby 2005).


Figure 1.23 Black coral at approximately 100 m depth
grown with the invasive snowflake coral Carijoa riisei.
credit: HURL (Hawaii Undersea Research Laboratory)
R. Grigg and S. Kahng.

Invasive Species
Invasions by non-indigenous marine species
have increased in the United States over the past
few decades due to increased shipping activities
that have incidentally transported species from
distant ports. The main vectors for transmission
of marine invasive species are ship ballast
water, hull fouling, and accidental or intentional
releases of exotic species from home aquariums
and scientific institutions. Because eradication
programs are rarely successful, preventing the

SJ ri LI I I J I I ^I I a '\ *wi '... Ji y LJ u I Ir
It has the potential to spread rapidly b:
budding and fragmentation of the mat
could promote rapid range expansion.

The rates of invasions and vectors c
transmission for invasive species ii

-I V y j I LI I-i IV-L IJI-IX \ I -I 11 1- III I 1 -,
to 90% of the black coral colonies were dead and
overgrown by the snowflake coral. This invasive
coral has been found overgrowing coral colonies
as deep as 120 m (Grigg 2003).

The invasive, colonial tunicate Didemnum sp.
was recently discovered on the northern edge
of Georges Bank, off New England. It is found
on hard substrates, and overgrows sessile and

,U biology and ecology of deep corals in
hives, general, impacts of invasive species
are difficult to measure.

Climate Change
Ocean Warming
The Intergovernmental Panel on Climate Change
(IPCC) was established in 1988 to assess
the risk of human-induced climate change, its
potential impacts, and options for adaptation and
mitigation. The IPCC 4th Assessment Report
(IPCC 2007a) concluded that since 1961, the
global ocean has absorbed more than 80% of
the heat added to the climate system. During
the period from 1961 to 2003, global ocean
temperature has risen by 0.1C from the surface
to a depth of 700 m (Bindoff et al. 2007), the
region where many deep corals are found.

Figure 1.24 The colonial tunicate Didemnum
sp. advances over a pebble and cobble habitat
on northern Georges Bank at a depth of 41 m
Photo credit: P. Valentine and D. Blackwood,
I I Q crnlnoinral Cn lm/


Increases in average temperature have affected
waters as deep as 3000 m. Eleven of the past 12
years (1995-2006) rank among the 12 warmest
years in the instrumental record of global surface
temperature since 1850 (IPCC 2007a). There
was significant bleaching of shallow-water
corals leading to mortality during this period,
especially in 1997/98 and in the Caribbean in
2005. The report concluded that it "is likely that
anthropogenic forcing has contributed to the
general warming observed in the upper several
hundred meters of the ocean during the latter
half of the 20th century" (IPCC 2007a) i.e., as a
result of the observed increase in anthropogenic
greenhouse gas concentrations. These projected
changes have been accompanied by observed
changes in ocean salinity and biogeochemistry
(Bindoff et al. 2007).

Model projections of future climate change
present a number of threat scenarios. Based
these scenarios the IPCC has concluded that
the most likely scenario likely that the Northern
Hemisphere thermohaline circulation meridionall
overturning circulation) of the Atlantic Ocean
will weaken during the 21st century (Joos et al.
1999; IPCC 2007a), but there is considerable
decadal variability in this circulation and data do
not support a coherent trend in the overturning
circulation (IPCC 2007a). Thermohaline
circulation is the major driving force behind
currents in the deep ocean. A weakening of
this process could reduce transport of food and
oxygen to deep coral communities and eventually
alter the structure of deep sea ecosystems. It
is unclear how these changes might affect deep

Ocean Acidification
The ocean acts as the largest net sink for CO2,
absorbing this gas from the atmosphere and
then storing carbon in the deep ocean. Since
pre-industrial times over half of the additional
CO2 attributed to human activities released
in the atmosphere has been absorbed by the
oceans (Sabine et al. 2004). The average pH
of the ocean has decreased by 0.1 units since
pre-industrial times, which represents a 30%
increase in the concentration of hydrogen ions
and is a geologically significant acidification
of the oceans (Caldeira and Wickett 2003;
IPCC 2007a). Oceanic uptake of CO2 drives
the carbonate system to lower pH and lower
saturation states with respect to the carbonate

minerals calcite and aragonite, the materials
used to form supporting skeletal structures
in many major groups of marine organisms,
including corals (Kleypas et al. 2006). While
the effects of ocean acidification on the marine
biosphere have yet to be documented in the field
(IPCC 2007b), numerous laboratory and large-
scale mesocosm studies have demonstrated
cause for concern. Model scenarios predict that
there will be a further decrease in pH of 0.5 units
by the year 2100 (Caldeira and Wickett 2005).
This change in ocean chemistry could reduce
the ability of corals to produce calcium carbonate
skeletons (calcification) and build reefs. Others
have predicted that ocean acidification as
a consequence of doubling preindustrial
atmospheric CO2 could decrease shallow coral
calcification rates by 10-30% (Gattuso et al. 1999;
Kleypas et al. 1999; Feely and Sabine 2004).
There is evidence that the rate of CO2 increase
in the deep ocean has been occurring at double
the pace of shallow waters, and therefore the
effect of ocean acidification on deep corals could
be significant (Bates et al. 2002).

There is a natural boundary in the oceans
known as the aragonitee saturation horizon"
below which organisms such as stony corals
cannot maintain calcium carbonate structures.
As CO2 levels increase, the aragonite saturation
horizon becomes shallower, severely limiting
the distribution of stony corals in certain parts
of the deep sea (Royal Society 2005; Guinotte
et al. 2006; Kleypas et al. 2006). Projected
increases in ocean acidity could result in severe
ecological changes for deep corals, and may
influence the marine food chain from carbonate-
based phytoplankton up to higher trophic levels
(Denman et al. 2007).

Other indirect effects of ocean acidification
on deep corals may involve changes in the
availability of nutrients and toxins. Changes
in pH could also cause a release of previously
bound metals from the sediment, increasing
the amount of metal toxins in the water column.
There is currently a need for long-term studies
on these effects in situ (Royal Society 2005).

Proposals have been made to capture CO2
emissions at the time of energy production
and inject it in the deep ocean, thus reducing
greenhouse gas emissions into the atmosphere.
Small-scale experiments and modeling suggest


Figure 1.25 Map showing the U.S. EEZ and areas of seven regional assessments.

that injected CO2 would be isolated from the
atmosphere for several hundred years (IPCC
2005). The long term results to the atmosphere
of large scale experiments are unclear, but these
actions could drastically change the chemistry
of the deep ocean. In the recent IPCC Special
Report (2005), methods detailing carbon dioxide
capture and storage were discussed. The
report warned that not enough is known about
the effects of excess CO2 on benthic marine
organisms. Few countries currently have
legal or regulatory frameworks for dealing with
environmental impacts of CO2 sequestration.

The cumulative impacts on deep coral
ecosystems of warming ocean temperatures and
ocean acidification due to climate change are
still unknown. They may be secondary stressors
on corals already impacted by other threats or

National Overview

In the chapters that follow, the authors draw
together current knowledge, including previously
unpublished data, on the distribution of deep


coral communities in seven broad regions of
the United States (Figure 1.25); the threats they
face; and current management efforts to address
these threats. The purpose of this section is to
provide a brief synthesis of some of the trends
found within and across the regions.

The U.S. exclusive economic zone (EEZ)
extends 200 nautical miles (370 km) offshore,
covering 11.7 million square kilometers in the
Pacific, Atlantic, and Arctic Oceans. This broad
geographic range includes a wide variety of
deep-water ecosystems, most of which have not
been explored. Of what is known, large regional
differences exist among the types of corals
present, theirassociated communities, the amount
of available regionally specific information,
and the methods applied to characterize and
understand deep coral communities. The threats
faced by deep coral communities differ regionally,
as do the management approaches that have
been adopted to address impacts from fisheries.
The research and exploration work conducted
over the past 20 to 30 years has helped pave
the way to understanding these ecosystems and
addressing management concerns, but it is only
a start.


Important deep coral communities have been
identified in every U.S. region. Most deep coral
groups, with the exception of pennatulaceans,
occur primarily on hard substrata, especially
near the continental shelf break, along the
continental slope, and on oceanic islands
and seamounts. The distribution of individual
species is determined in part by major currents,
and their interaction with local geomorphology of
seamounts and coastal shelves.

Currently, it is impossible to ascertain the overall
extent of deep coral communities, much less
their condition or conservation status in U.S.
waters, because so many of the deeper areas
these communities inhabit have been explored
incompletely or have not been explored at all.
There is also very little information on what most
continental shelf habitats may have looked like
before there was extensive trawling. Therefore,
the following apparent trends in distribution
should be viewed with caution.


The major oceanographic influences on the
U.S. Pacific Coast are the Alaska and California
Currents, which are formed when the eastward-
flowing North Pacific Current bifurcates near
Vancouver Island. The counterclockwise Alaska
Current continues along the southern edge of
the Aleutian Islands as the Alaska Stream, and
a parallel low salinity Alaska Coastal Current
flows close to the coast from British Columbia
to Unimak Pass, and into the Bering Sea. The
California Current moves south, transporting cold
northern waters along the Washington, Oregon,
and California coasts. The northeast Pacific is
characterized by a relatively narrow continental
shelf with active tectonic and volcanic processes,
creating a complex bathymetry of canyons
and other features that support rich benthic
communities. Prevailing spring and summer
winds cause upwelling of deep nutrient-rich
waters that influence production over the shelf.

The widespread U.S. oceanic islands and
associated seamount chains in the tropical
central and western Pacific are volcanic in origin
and subject to various major currents. Localized
flow around pinnacles, seamounts, and oceanic
islands likely has the largest effect on the local
abundance of deep corals. Seamounts, in any
ocean basin, obstruct ocean currents and by
doing so create eddies and local upwelling, form
closed circulation patterns called Taylor columns,
and enhance local production (Boehlert and
Genin 1987). As a consequence seamounts
possess both hard substrate and high flow,
ideal conditions for the development of deep
coral communities. The presence of seamounts
and oceanic islands can have an effect on local
production. In comparison to adjacent oceanic
water masses, the waters around seamounts
have been noted to have higher nutrient
and chlorophyll concentrations, and higher
zooplankton, ichthyoplankton, and micronekton

A major distinction between the North Pacific and
the North Atlantic coral communities was thought
to be the absence of stony coral bioherms or
reefs in the deep waters of the North Pacific
(Freiwald et al. 2004). Only isolated records of
the stony coral Lophelia pertusa and other reef-
builders had been reported. Stony corals accrete
calcium carbonate in the form of aragonite, and


accretion rates must exceed dissolution rates
for reef structures to be built. Guinotte et al.
(2006) noted this apparent absence of reported
stony coral bioherms in the North Pacific, and
hypothesized that it might reflect the shallow
depth of the aragonite saturation horizon in the
North Pacific (50-600 m). This horizon reaches
depths of more than 2,000 m in the North Atlantic,
where deep stony coral reefs have been best
studied. The absence of deep coral reefs was
recently called into question with the discovery
of patchy, low-lying accumulations of live and
dead L. pertusa off the coast of Washington
State (Chapter 3; Hyland et al. 2005; Brancato
et al. 2007). Because of the lack of massive
structures in the Pacific similar to those seen
in the Atlantic, it is not clear from these initial
reports whether these lower-lying accumulations
might be classified as reefs or bioherms.

Alaska Region: The U.S. EEZ around Alaska
includes the Gulf of Alaska, Aleutian Islands, and
easternBering Sea in the Pacific, and the Chukchi
and Beaufort Seas in the Arctic. Deep corals
are an important structural component of the
first three of these Alaskan marine ecosystems
(Chapter 2). Gorgonian deep corals reach their
highest diversity in the Aleutian Islands, often
forming structurally complex "coral gardens"
with stylasterid corals, sponges, and other
sedentary taxa. Gorgonians are also the most
important structure-forming corals in the Gulf
of Alaska, with species of the genus Primnoa
reaching 5-7 m in size, while the Bering Sea
has dense aggregations of soft corals and sea
pens on the shelf and slope, respectively. The
region is relatively depauperate in scleractinian
corals, which occur as solitary cups and do not
form true coral reefs. Most information on the
distribution of deep corals comes from NOAA
trawl surveys, supplemented more recently
by NOAA submersible and remotely operated
vehicle (ROV) studies conducted on the shelf
and slope of the Aleutian Islands and Gulf of
Alaska, and on seamounts in the Gulf. Currently
there is very little information on deep corals in
the Arctic Ocean.

The region supports some of the most important
groundfish and crab fisheries in the world. It also
appears to have the best-developed information
on the association of fish species with many of
these deep octocoral resources (Chapter 2;
Heifetz 2002; Krieger and Wing 2002; Stone

2006). As some of the same coral and fish
species (e.g., rockfishes) also occur along the
West Coast region, it is possible that some of
these fish/coral associations may also occur

U.S. West Coast Region: The Pacific waters off
the Washington, Oregon, and California coasts
are part of the California Current Large Marine
Ecosystem (LME). The deep coral communities
in this region are similar to those farther north
along the Pacific coasts of British Columbia
(Etnoyer and Morgan 2005) and Alaska (Chapter
2). As in Alaska, understanding of the spatial
extent of these communities has benefited from
relatively extensive NOAA trawl survey catch
records, supplemented by museum collections
and in situ observations.

Gorgonians are the most abundant and diverse
structure-forming deep corals along the West
Coast (Figure 1.26). As elsewhere, these
appear especially associated with hard, exposed
substrata and steeper slopes. There appear to
be biogeographic differences in the distributions
of certain deep coral groups within the region.
Gorgonians appear to be most abundant south of
Point Conception and north of Cape Mendocino
(Chapter 3; Etnoyer and Morgan 2003). Black
corals (Figure 1.27) appear abundant between
Cape Mendocino and Canada.

U.S. Pacific Islands Region: The U.S. Pacific
Islands represent diverse oceanic archipelagos
scattered across wide areas of the Pacific and
encompassing several different biogeographic
regions. They do not have continental shelves
or slopes, but represent emergent and non-
emergent seamounts many highly isolated
from other areas. Aside from the Hawaiian
Archipelago, almost nothing is known of the
deep coral resources in the U.S. Pacific Islands.
The first submersible explorations of American
Samoa and the U.S. Line Islands were begun in
2005, and surveys of additional areas in the U.S.
Pacific are needed.

Octocorals and black corals are the principal
structure-forming species on deep Hawaiian
slopes and seamounts (Chapter 4). Taxonomic
surveys of deeper water scleractinians in
Hawaiian waters have been reported by
Vaughan (1907) and Cairns (1984, 2006).
While the Hawaiian Archipelago shares some


Figure 1.26 Paragorgia sp. crown a ridge on the Davidson Seamount. Photo credit: The Davidson Seamount
Expedition, MBARI, and NOAA-OE.

species with Alaska and the West Coast, it
is likely that it has a relatively high degree
of endemism; rates of endemism have been
estimated at 29% (Maragos et al. 2004) and 21%
(Cairns 2006) for the shallow-water and deep-
water scleractinian coral fauna, respectively.
Paradoxically, understanding of the unique deep
coral assemblages in Hawaii has benefited
from information gathered in association with
commercial harvests of deep corals including
gold (Gerardia sp.) and pink (Corallium spp.)
precious corals and the shallower black coral
(Antipathes spp.). Monitoring in support of
management has provided perhaps the most
extensive studies of growth and recruitment
rates for any deep coral taxa.


The Gulf Stream is the dominant oceanographic
feature influencing much of the U.S. Atlantic.
It originates in the Caribbean, flows through
a loop current in the eastern Gulf of Mexico,

exits through the Florida Straits, and moves
northward along the U.S. East Coast. Its depth
extends to areas where it may influence deep
coral distribution. Though the Gulf Stream is
diverted eastward at Cape Hatteras, it still has
great influence in northeast regional waters,
interacting with a southwest flow of coastal
waters and contributing to gyres and complex
circulation patterns.

U.S. Northeast Region: The Northeast U.S.
Continental Shelf Large Marine Ecosystem6
(LME) and associated continental slope extend
along the Atlantic coast from the Gulf of Maine
to Cape Hatteras, with a number of seamounts
occurring in the New England area. This region
has among the longest histories of both deep sea
scientific research and extensive trawl fisheries.
Understanding of coral resources in the region
has benefited from the work of Theroux and
Wigley (1998) and especially Watling et al.
'For more information on Large Marine Ecosystems and
their designation visit www.edc.uri.edu/Ime


Figure 1.27 Recently discovered Christmas tree coral, Antipathes dendrochristos. Photographed from Delta
submersible during surveys of deepwater rocky banks off southern California. Photo credit: M. Amend.

(2003) in mapping the reported occurrences of
major deep coral species. Gorgonians represent
the predominant structure-forming deep coral
taxa in this region, and they appear to be most
numerous on hard substrates associated with
canyons along the shelf and Georges Bank
slopes, and on the New England Seamount chain
(Chapter 5; Auster et al. 2005). The principal
species recorded in this region have also been
recorded in Canadian waters (Gass and Willison
2005). Although L. pertusa has been infrequently
reported from waters off the northeastern U.S.,
no major reef-like formations are known to
exist. Such formations are common south of
Cape Hatteras (Chapter 6), and known from
at least one location in Atlantic Canada the
Stone Fence reef at the mouth of the Laurentian
Channel (Gass and Willison 2005).

Significant concentrations of gorgonians have

been recorded from Oceanographer and Lydonia
Canyons on Georges Bank and from Baltimore
and Norfolk Canyons further south. It is not clear,
however, to what extent these reports reflect only
areas where studies have been conducted. It is
possible that significant additional information
on coral distributions can be mined from NOAA
trawl surveys conducted in this region. Recent
expeditions to the New England Seamounts
(Chapter 5; NOAA Ocean Exploration 2005,
North Atlantic Stepping Stones) have also
revealed unique assemblages of deep corals on
these seamounts.

U.S. Southeast Region: Based on regions
surveyed within U.S. waters, deep-water
scleractinian coral reefs probably reach their
greatest abundance and development in the
Atlantic, south of Cape Hatteras (Chapter 6).
Information from this region is primarily derived


Figure 1.28 Multi-colored gorgonians with whip coral in background. East Bank, Flower Garden Banks NMS
Photo credit: NURC/UNCW and NOAA/FGBNMS

from research submersible studies at isolated
locations and soundings that have revealed
potential coral banks. L. pertusa is the major
structural component of reefs on the continental
slope and Blake Plateau from North Carolina to
Florida. It provides habitat for a well developed
faunal community that appears to differ from the
surrounding non-reef habitats (Chapter 6). This
region is influenced by the Gulf Stream, which
may contribute to biogeographic linkages between
the southeast U.S. and better studied northeast
Atlantic Lophelia ecosystems. The world's only
known Oculina varicosa reefs are found in 70-
100 m depths off east-central Florida. Because
of their shallow depth, and occurrence on the
continental shelf, they may differ from other deep
coral reefs in structure, function, or composition
of associated organisms. The shallow depth
range of these reefs has facilitated a more
comprehensive understanding of the ecology of
the corals; the role of the reefs as essential fish
habitat; and the impacts of trawl fishing on these
resources (Chapter 6; Koenig 2001; Reed 2002b;
Koenig et al. 2005). Gorgonians are common in
the region, but in comparison to the Northeast
and West Coasts, much less is known (or at
least less information has been systematically
collated) concerning the region's octocoral and
black coral resources.

U.S. Gulf of Mexico Region: The northern
Gulf of Mexico is home to major L. pertusa
reefs, though their structure appears to differ

from that observed in the southeast U.S.
(Chapter 7), growing primarily on carbonate
and clay substrates rather than mounds of dead
coral. Despite extensive environmental studies
associated with oil and gas development in
the Gulf, knowledge of the distribution of deep
coral reefs is limited to a handful of sites where
targeted studies have been conducted. Each
of the areas, from Pourtales Terrace in the
Florida Straits, to sites in the northwestern Gulf
of Mexico, represent unique habitat types. As
in the Southeast, little information is available
concerning the distribution of gorgonian and
black coral resources in this region (Figure 1.28).
Cairns and Bayer (2002) identify several species
of the structure-forming primnoid Callogorgia
occurring throughout the Gulf. Of these, the
endemic gorgonian C. americana delta is
known to provide nursery habitat for catsharks
(Etnoyer and Warrenchuk in press). Recent
ROV surveys focused on the reefs and banks
of the northwestern Gulf of Mexico at depths
ranging from 50 m to 150 m have increased
our knowledge of the distribution of deepwater
biological communities, including antipatharians,
gorgonians, and sponges (Figure 1.29). The
communities are more widespread and densely
populated than reported thus far. These studies
are ongoing, and being led by the Flower Garden
Banks National Marine Sanctuary, (E. Hickerson
and G.P. Schmahl, pers. comm.).


Figure 1.29 An example of deepwater habitat at the Flower Garden Banks NMS, typical of the northwest Gulf
of Mexico habitats. Image includes octocorals, antipatharians, echinoderms, sponges, soft corals, and deep-
water fishes. Photo credit: FGBNMS and NURC/UNCW.

U.S. Caribbean Region: The U.S. Caribbean,
including the waters surrounding Puerto Rico,
the U.S. Virgin Islands, and Navassa Island,
represents a small part of the larger Caribbean
LME. It has not been well studied with respect to
deep corals, and the primary information comes
from scientific collections (e.g., Cairns 1979)
- most from other areas of the wider Caribbean
(Chapter 8). The most extensive occurrence of
deep coral mounds reported in the Caribbean
is found on the northern slope of Little Bahama
Bank at depths between 1,000 and 1,300 m
(Chapter 8). Lithoherms have been documented
in the Florida Straits, and deep coral banks are
known to occur off Colombia's Caribbean shelf.
There is some indication that the diversity of
certain deep-water structure-forming taxa (e.g.,
gorgonians) may be higher in the Caribbean than
in more temperate North Atlantic waters. In U.S.
waters, limited ROV and submersible studies
have been conducted off Navassa Island and
Puerto Rico, revealing scleractinian, black, and

gorgonian corals, but distributions have not been
rigorously documented.


Summary of Threats to Deep Coral
Communities in U.S. Regions

In the chapters that follow, the authors identify
key threats to deep coral communities in each
region. The perceived level of each of these
key threats is summarized in Table 1.3. Each
region has different intensities of trawl fishing
and different levels of information on the actual
impacts of such fisheries. However, based on
the best available data, disturbance to deep
coral communities from bottom-tending fishing
gear, especially bottom trawl gear, has been
identified as the major concern in most regions
where such fishing is allowed. Similar findings
have also been reported from elsewhere around


Table 1.3. Summary of perceived levels of currer
Applicable (i.e., this threat is prohibited or does n
on the information provided by the regional chapi
Note: These perceived threat levels reflect only th
if unmitigated, to damage deep coral communities.
of each stressor, which will likely vary widely with
is incompletely known, there is uncertainty ove
management steps have been taken to mitigate
impacts of bottom fishing gear through gear mo(
and management procedures are in place to miti
where they occur on the outer continental shelf.

West Pi
Threats Alaska Coast IslI
Bottom trawl
fishing impacts I
Other bottom Low Low -
fishing impacts Medium Medium L
Deep coral harvest NA NA
Oil and gas
development Low Low I
Cable deployment Low Low Unl
Sand and gravel
mining Low NA I
Invasive species Unknown Unknown
Climate change Unknown Unknown Unk

the world (e.g., Rogers 1999; Koslow et al. 21
Hall-Spencer et al. 2001; Fossa et al. 21
Roberts 2002; Grehan et al. 2005, Wheeler E
2005). Harvest of black and precious cora
Hawaii has been identified as a moderate thi
but it is conducted in a very selective mar
and its overall impact is minor compared to t
fishing in other regions. Hawaii is also the
jurisdiction that has specifically identified a cur
threat to deep corals from an invasive spec
Oil and gas exploration and development in
Gulf of Mexico, where it is increasingly mo
into deeper waters, is the only non-fishing d
anthropogenic stressor that poses a mode
threat to deep coral communities. Pote
impacts from climate change (Including oc
acidification) are largely unknown.

U.S. Management of Deep Coral Ecosyste
in an International Context

Recent interest in deep coral ecosystems
galvanized the public and triggered conserve
and management action in the United States
around the world. In recent years, conserve

treats to deep coral communities for U.S. regions. NA =
occur anywhere within that region). Threat levels are be
occurrence of these stressors in a region, and their potei
ey might encounter. They do not indicate the actual imp
nd among regions. Since the location of deep coral hab
heir degree of overlap with human activities. Substa
'eats. For example, significant actions to minimize adv
nations and gear closures have been taken in each re(
:e potential impacts of oil and gas development and mi

ic Gulf of
js Northeast Southeast Mexico Caribbe
Low -
Medium NA
Low Low Low -
Medium Medium Medium Low

wn Low Low Low Unkno,

Low Low Low NA
Unknown Unknown Unknown Unkno,
wn Unknown Unknown Unknown Unkno,

); actions have been taken shortly after disco,
2; of vulnerable deep coral habitats. Internation
II. this includes new marine protected ar
n established to protect deep coral commun
t, in the northeast and northwest Atlantic, in
r, Canadian Pacific, and on seamounts in Austi
vl and New Zealand. Internationally, a se
ly of United Nations General Assembly (UN
it resolutions has addressed the impacts of fisl
s. on vulnerable marine ecosystems in internatic
e waters, with specific reference to seamou
g hydrothermal vents, and cold-water corals.
:t international effort culminated in the Decen
:e 2006 UNGA Sustainable Fisheries Resolu
al (A/61/105), which calls upon states and regic
n fisheries management organizations (RFM
to ensure the sustainable management of
stocks and protection of vulnerable ma
s ecosystems including seamounts, hydrother
vents, and cold-water corals from destruc
fishing practices by December 31, 2008.
n In the United States, on October 6, 2(
d President Bush put forth a memorandurr
n promote sustainable fishing and end destruc

fishing practices. This memo called upon the
Departments of State and Commerce to work with
other countries and international organizations
to eliminate destructive fishing practices; work
with RFMOs to establish regulations to promote
sustainable fishing; develop new RFMOs to
protect ecosystems where they do not currently
exist; work with other countries to determine
which vulnerable marine ecosystems might be
at risk; and combat illegal, unregulated, and
unreported fishing. The memorandum defines
"destructive fishing practices" as "practices that
destroy the long-term natural productivity of fish
stocks or habitats such as seamounts, corals
and sponge fields for short term gain."

In addition to addressing the effects of fishing
on deep coral habitats, other multilateral
environmental fora have addressed deep-sea
genetic resources and the impacts of trade.
All black, hydrozoan (e.g., stylasterid), and
stony corals are included in Appendix II of the
Convention on International Trade in Endangered
Species of Fauna and Flora (CITES). These
listings still allow trade under permit, but they are
designed to ensure the harvest and trade is legal
and non-detrimental to wild populations.

U.S. National Framework for Management of
Deep Coral Ecosystems

In the U.S., management of deep coral resources
has been hampered by a lack of information
on the distribution, life history, and ecological
role of these organisms. Deep corals were
not specifically included in legislation prior to
the Magnuson-Stevens Fishery Conservation
and Management Reauthorization Act of 2006
(PL. 109-479). Currently, no deep-water coral
species are listed as endangered or threatened
under the Endangered Species Act, nor are
any presently under consideration for listing,
although 0. varicosa has been identified as
a "species of concern."7 In a number of U.S.
regions, significant management measures are
now being undertaken and these efforts will be
discussed in the regional chapters in detail but
are summarized in this National Overview. As
fisheries expand into deeper waters (Roberts
7"Species of concern" are species about which NMFS has
some concerns regarding status and threats, but for which
insufficient information is available to indicate a need to list
the species under the Endangered Species Act. http://www.


2002), and oil and gas exploration and
development activities move to deeper areas of
the continental slope, precautionary measures
should be taken to preserve the fragile biota that
exist in those areas.

Most deep corals occur in the U.S. EEZ beyond
the jurisdiction of individual states. Fisheries
in the EEZ are managed by NOAA's National
Marine Fisheries Service (NMFS) under fishery
management plans (FMPs) prepared by eight
regional Fishery Management Councils (FMCs)
and approved by NMFS in accordance with
the Magnuson-Stevens Fishery Conservation
and Management Act (16 U.S.C. 1801 et seq.).
These eight Council regions align closely with
the boundaries of the regional chapters in
this report. As each Council region includes
different fisheries and has developed FMPs
independently, approaches to deep coral
conservation also vary. To date, management
approaches by the Councils to reduce fishery
impacts on deep corals (Table 1.4) have primarily
relied upon either treating the corals themselves
as a managed species (South Atlantic and
Western Pacific Councils) or protecting habitats
identified as essential fish habitat (EFH) for
managed species that may contain deep corals
(South Atlantic, North Pacific, Pacific, and New
England Councils). In the New England and
Mid-Atlantic Council regions, where deep corals
have not been specifically identified as EFH,
the scope for using these provisions to protect
coral habitat may be more limited. Councils are
also mandated to minimize bycatch to the extent
practicable, but none have used this provision
directly to regulate bycatch of deep corals.

On January 12, 2007, the Magnuson-Stevens
Fishery Conservation and Management
Reauthorization Act of 2006 (PL. 109-479) was
enacted and included the "Deep Sea Coral
Research and Technology Program." The Act
calls on NOAA to: 1) identify existing research
on, and known locations of, deep-sea corals
and submit such information to the appropriate
Councils; 2) locate and map locations of deep-
sea corals and submit such information to the
Councils; 3) monitor activity in locations where
deep-sea corals are known or likely to occur,
based on best scientific information available,
including through underwater or remote sensing
technologies, and submit such information to
the appropriate Councils; 4) conduct research,


Table 1.4. Regional fishery management council management actions affecting deep coral habitat.

Yes Groundfish No Yes Yes
Yes Groundfish No Yes Yes
Yes Precious
No e o Yes Not Applicable
Limited Lydonia &
No No Oceanographer Canyons No
and New England Groundfish
Habitat Closure Areas
No No No No
Yes Used to
Yes- Snapper e -Us Limited Oculina Banks
Grouper Complex HAPC
Yes Not yet
No applied to deep No No
Yes Not yet
No applied to deep No Not Applicable

including cooperative research with fishing
industry participants, on deep sea corals and
related species, and on survey methods; 5)
develop technologies or methods designed to
assist fishing industry participants in reducing
interactions between fishing gear and deep-
sea corals; and 6) prioritize program activities
in areas where deep-sea corals are known to
occur, and in areas where scientific modeling or
other methods predict deep sea corals are likely
to be present. The first biennial report on the
progress and significant findings of the "Deep
Sea Coral Research and Technology Program"
is due to Congress by January 12, 2008. The
Act also provides new discretionary authority for
fishery management plans to designate zones
where deep-sea corals are identified through the
program to protect deep sea corals from physical
damage from fishing gear or to prevent loss or
damage to such fishing gear from interactions
with deep-sea corals, after considering long-term
sustainable uses of fishery resources in such

In addition to the Councils, NOAA's National
Marine Sanctuary Program has responsibilities
for protection and management of natural
resources, and a number of sanctuaries contain

deep coral resources. The goals of the National
Marine Sanctuaries Act (16 U.S.C. 1431 et
seq.) include maintaining the natural biological
communities in the national marine sanctuaries,
protecting, and, where appropriate, restoring,
and enhancing natural habitats, populations
and ecological processes. New oil and gas
development is currently prohibited in all national
marine sanctuaries, although leases in place
before sanctuary designation (e.g., Channel
Islands National Marine Sanctuary) are allowed
to continue. Roughly half of the national marine
sanctuaries have regulations that prohibit
activities (some specific to fishing) that could
damage deep coral communities. Further, a
number of sites have recently taken specific
actions to characterize and protect deep coral
communities, in particular Flower Garden Banks,
Olympic Coast, Monterey Bay and, indirectly,
Channel Islands, with new marine protected area
designations. These sanctuaries have been
extremely active with deep coral community
characterization Monterey Bay National
Marine Sanctuary at Davidson Seamount
and other sites, the Flower Gardens on outer
continental shelf banks of the Gulf of Mexico,
and Olympic Coast in the Pacific Northwest
(Figure 1.30). Deep coral communities are


Figure 1.30 Rockfish take refuge among a primnoid octocoral in Olympic Coast National
Marine Sanctuary. Photo credit: OCNMS/NOAA

found in the Papahanaumokuakea Marine
National Monument (see Chapter 4). The
National Marine Sanctuaries Act may provide
more comprehensive protection in these areas
from collecting, development, discharges, and
other human activities that disturb benthic
habitats. Deep coral communities may also
occur in certain National Parks and National
Wildlife Refuges, especially in Alaska and the
Pacific remote island areas.

Mineral resource exploration and extraction
activities, including oil and gas exploration in
federal waters, are managed by the MMS within
the U.S. Department of the Interior. The MMS
regulates the impact of mineral resource activities
on the environment through an Environmental

Studies Program and an Environmental
Assessment Program. These programs provide
scientific and technical information to support
decisions and monitor environmental impacts
of exploration, development, and production
of mineral resources. MMS established the
Rigs-to-Reefs program to explore the use of
decommissioned oil platforms as hard substrate
for settlement and growth of corals and other
sedentary marine organisms.

Regional Management Actions in the
U.S. Pacific

Acknowledgement of the potential impacts of trawl
and dredge fisheries to deep coral communities
and other biogenic habitat has led the regional


Fishery Management Councils in the Pacifi
propose historic protective measures lim
bottom-trawling in areas that might contain c

The U.S. West Coast and Alaska have
extensive history of fisheries using boti
contact gear, including bottom trawling for Pa
cod, hake and rockfishes; bottom-set long
for fish; and individual traps and multiple t
lines for crab in Alaska. Alaska is currently hi
port to the largest fleet of U.S. bottom traw
The importance of these bottom-trawl fisher
has been a major factor in the develop
of NMFS trawl surveys. These surveys I
provided the broadest scale information on
distribution and abundance of deep coral
these two regions (Chapters 2 and 3).

The North Pacific Fishery Management Cot
has taken a number of important steps that rec
the impact of fisheries on essential fish habit
the EEZ around Alaska. Beginning in 1998,
Council prohibited trawling in the eastern GL
Alaska and southeast Alaskan waters with
180,400 km2 area as part of a license-limits
program. The measure was originally propc
in 1991 over conservation concerns for rod
stocks to protect seafloor habitat from long-1
disturbance from trawling. In 2000 the Cot
established the 10.6 km2 Sitka Pinnacles Me
Reserve in the Gulf of Alaska and prohibit
bottom-fish gear types (except pelagic troll
for salmon) in the reserve. These pinna
consist of two large volcanic cones that ris
within 40 and 70 m of the ocean surface,
provide a variety of high-relief habitats colon
by the deep coral Primnoa sp., anemones,
other organisms. Aggregations of lingcod
several juvenile and adult rockfish species
associated with the pinnacles.

Recently, the North Pacific and Pacific Fisl
Management Councils each took historic st
recommending to "freeze the footprint" of boti
trawling within their respective jurisdiction
order to protect EFH. In 2006, NMFS apprc
a number of North Pacific Council recommer
EFH closures in the Aleutian Islands and
of Alaska. Many of these areas are kn
to contain important deep coral and spc
habitats. More than 950,000 km2 along
remote Aleutian Islands were closed to bol
trawling targeting areas that had not

:o received extensive trawling, with 377 km2 of "c
g gardens" closed to all bottom-tending fisl
al gear. Additionally, 7,155 km2 in Gulf of Ale
Slope Habitat Conservation Areas were clc
to bottom trawling and 18,278 km2 of Ale
n seamounts and 46 km2 of Primnoa coral ar
i- in the Gulf of Alaska were closed to all bott
ic tending fishing gear. In June 2007, the N
Gs Pacific Fishery Management Council adol
- additional measures to conserve benthic
e habitat in the Bering Sea. These measure
s. approved, NMFS would prohibit bottom trav
is over an additional area of more than 450,
it km2.
e The Pacific Fishery Management Counci
n responsible for developing FMPs for fisheries
the coasts of California, Oregon, and Washing
Within the past three to six years, comme
:il fishing has been prohibited or significe
:e curtailed within the Cowcod and Rocl
n Conservation Areas. While these restrict
e were not designed to address impacts on d
)f corals, they are likely to protect some deep c
a habitats. Beginning in 2000, the Council
n prohibited footrope trawls (footrope=weigl
d edge of trawl that impacts seafloor) gre
h than 8 inches on most of the continental sl
n effectively making many complex, rocky hab
:il that are home to deep corals inaccessible
e trawlers.
ar In 2006, NMFS approved a plan that ident
is and described EFH for Pacific groundfish
:o prohibited bottom trawling in 336,700 km
d habitat off the West Coast of the U.S.
d represents over 42 percent of the EEZ
d Washington, Oregon, and California, inclui
d areas that may contain deep coral and spo
-e habitats. Selected areas with known d
coral resources (e.g., Davidson Seamount)
protected from all bottom-contact gear.
s, Unlike most other areas of the United States,
i- Insular Pacific has no history of domestic bott
n trawl fisheries. The Western Pacific Fisi
d Management Council manages the fisherie
d federal waters around the Territory of Amer
ilf Samoa, Territory of Guam, State of Ha\
'n Commonwealth of the Northern Mariana Islai
e and other U.S. Pacific island possessions. It
e the oldest and most comprehensive restrict
n designed to protect coral and other biog
et habitat from adverse impacts of fishing g


983, the Council prohibited the use of trawl G
r, bottom-set longlines, and bottom-set fc
ets all identified as threats to deep corals di
:hin all waters in their region of the U.S. EEZ. lir
Action was taken, in part, in response to g9
served impacts of foreign trawl fisheries on n,
counts (e.g., Hancock Seamount) before the tl
aration of the U.S. EEZ. th
383, the Western Pacific Fishery Management
ncil also developed a Precious Corals FMP. T
coral beds included in the FMP contain h,
eral deep coral species (Chapter 4). Under C
1983 FMP and its amendments, NMFS F
iblished quotas and minimum legal sizes for re
'est of pink, black, gold, and bamboo coral, A
, in 2002, prohibited the use of non-selective re
r. Currently, only the black coral fishery oi
in Hawaii State waters is active, and the e,
ncil and State are in the process of revising tr
iagement plans that incorporate the recent C
acts of the invasive soft coral Carijoa riisei. th
rional Management Actions C
ie U.S. Atlantic ki
Northeast U.S. region has the longest history re
najor trawl and scallop dredge fisheries in cc
United States and its bottom-trawl fishery a:
second in size only to Alaska's. As a result, P
;h of the continental shelf has been heavily
'led or dredged. In addition to trawling, there T
active fisheries using bottom-set longlines, b,
ets, and pots and traps, some extending into in
slope and canyon habitats that are known th
p coral habitat. (r
New England Fishery Management Council in
ages fisheries off the coast of Maine, New ui
ipshire, Massachusetts, Rhode Island, rr
Connecticut. The Mid-Atlantic Fishery K
iagement Council is responsible for C
lagement of fisheries in federal waters off the tr
sts of New York, New Jersey, Pennsylvania, T
aware, Maryland, Virginia, and North e:
olina. b'
two Councils oversee significant trawl p,
dredge fisheries with potential impacts A
deep coral habitats. In 2005, in order to C
mize adverse impacts to EFH, NMFS rr
roved Council-recommended closures ol
Oceanographer and Lydonia Canyons w
)roximately 400 km2, on the southern flank of

rges Bank), to bottom trawling and gillnetting
monkfish. These canyons are areas of known
) coral communities. Also approved were
s on the size of the bottom-trawling roller
r and rockhopper gear on the footrope of the
to no more than six inches in diameter in
submarine canyon areas off the shores of
mid-Atlantic states known as the "southern
agement area" of the monkfish fishery. In

South Atlantic Fishery Management Council
jurisdiction over FMPs in the EEZ off North
)lina, South Carolina, Georgia, and eastern
da to Key West (note: North Carolina is
esented in both the Mid-Atlantic and South
itic Councils). Trawling in the south Atlantic
on is primarily limited to shrimp trawling
he continental shelf. The Council was an
/ leader in addressing the threats of bottom
ling to deep coral communities. In 1984, the
ncil established the 315 km2 Oculina HAPC,
Norld's first protection granted specifically to
,ep coral habitat. In 2000 the South Atlantic
ncil expanded the Oculina HAPC to 1,043
The South Atlantic Council is currently
)wing and evaluating options for gear
ilations and four new HAPCs containing deep
I habitats, including two very large areas,
'art of a Comprehensive Fishery Ecosystem
I Amendment (Chapter 6).

habitats to cons

narily rock shrimp and calico scallop). Despite
/ protection, enforcement difficulties resulted
)ntinued destruction through illegal fisheries
recent requirements for use of vessel
itoring systems and enhanced enforcement.
nig et al. (2005) estimated that 90% of
lina coral banks had been damaged by
ling by 2001 and only 10% remained intact.
is perhaps the clearest U.S. example of the
nsive damage to deep coral communities
:rawling. Oculina varicosa was identified
JMFS in 1991 as a "candidate species" for
ntial listing under the Endangered Species
based on well-documented declines in the
lina coral banks areas due to damage from
hanical fishing gear, coupled with a lack of
served recruitment. In 2000 this designation
revised to "species of concern."


In addition to addressing fishing impacts, the
State of Florida has been proactive in the
management of potential new threats. Liquid
natural gas ports and pipelines are being
proposed that could impact deep coral habitats.
Florida is also a major hub for fiber optic cable
connections throughout the Caribbean. The
State of Florida has been a leader in developing
incentives for companies to locate cables in less
environmentally sensitive corridors.

In the central and western areas of the U.S.
Gulf of Mexico, concerns over potential damage
from fisheries are overshadowed by issues of
oil and gas exploration and development. With
over 4,000 active leases in depths inhabited
by Lophelia coral (deeper than 300 m), there is
potential for adverse interactions. A strategy,
developed in 2003 to address post-lease National
Environmental Policy Act compliance in deeper
waters (>400 m), requires lessees and operators
to submit an exploration plan for an ROV survey
of well sites. The plan requires a visual survey
of the seafloor in the vicinity of the well before
and then immediately after drilling activities
to ensure that drilling activities do not have
impacts on local benthic fauna. Almost half of
the deep-water lease sites have been thoroughly
surveyed with ROVs to document the biological
communities found in these areas (MMS 2003).
Along the continental shelf of the northwestern
Gulf of Mexico, dozens of reefs and banks
harbor deepwater communities of antipatharians,
gorgonians, and sponges, in depths from 50 m to
150 m. The MMS has provided protection from
direct impacts from oil and gas activities through
the topographic features stipulation, which
places "no-activity" zones and other regulatory
zones around these biologically sensitive areas.
These zones will be re-evaluated based on newly
acquired bathymetry.

The Gulf of Mexico Fishery Management Council
has jurisdiction over FMPs in the federal waters
off Texas, Louisiana, Mississippi, Alabama, and
the west coast of Florida. The primary fishing
impacts of concern to deep corals in the Gulf of
Mexico revolve around limited deep-water trawl
fisheries for royal red shrimp. The Council has a
Coral FMP and has protected several shallow-
water coral banks, but has not yet identified
deep coral habitat areas of particular concern.
Fishing restrictions through the Coral EFH of the
HAPC designation prohibit bottom longlining,

bottom trawling, buoy gear, dredge, pot, or trap
and bottom anchoring by fishing vessels at West
and East Flower Garden Banks, Stetson Bank,
McGrail Bank, and an area of Pulley's Ridge.
Other NW GOM HAPC's that do not carry any
regulations are in place at 29 Fathom, MacNeil,
Rezak, Sidner, Rankin, Bright, Geyer, Bouma,
Sonnier, Alderdice, and Jakkula Banks. The
Council recently asked its Coral Scientific and
Statistical Committee to develop a research
approach to identify locations of deep corals in
the Gulf.

Although not expressly prohibited, there is no
history of trawl fisheries in the U.S. Caribbean.
Fish traps are commonly used in shallower
waters, but deeper areas are not targeted.
The Caribbean Fishery Management Council
has jurisdiction over FMPs in federal waters
surrounding the Commonwealth of Puerto
Rico and the United States Virgin Islands. The
Caribbean Council has a Corals and Reef
Associated Invertebrates FMP, but, like the
Gulf of Mexico Council, it has not proposed
management measures that would specifically
identify deep coral areas. Navassa Island,
claimed by both the United States and Haiti,
is administered by the United States Fish and
Wildlife Service, which manages the Navassa
Island National Wildlife Refuge.


The authors of each of the regional chapters
have identified research priorities for their region.
The following research priorities are common to
several or all regions, or areas of research that
transcend regional interests and boundaries
and would contribute directly to improved
management. Most of these priorities address
information related to identifying locations of deep
coral communities and the status and trends of
deep corals and their associated communities,
and do not represent a comprehensive list of
scientific research needs (see also McDonough
and Puglise 2003; Puglise et al. 2005). In situ
research on deep coral communities requires
the use of specialized types of underwater


Habitat Mapping and Characterization

The highest priority in every region is to locate,
map, characterize, and conduct a baseline
assessment of deep coral habitats. The
location of deep coral habitats is not well known,
making it difficult if not impossible to adequately
protect these habitats and manage associated
resources. Acoustic multibeam bathymetry maps
and associated backscatter imagery at depths
between 200 and 2,000 m on continental slopes
and seamounts are basic tools for determining the
potential distribution of deep coral communities.
Bathymetric maps of underwater topography can
identify areas of potential coral habitat, based
on slope or other physical features (Morgan et
al. 2006), which can then be prioritized for more
detailed study. Multibeam backscatter imagery
provides clues as to substrate hardness. With
the exception of sea pens (pennatulaceans),
the major structure-forming deep corals are
dependent upon exposed hard substrata for
attachment. Though in certain cases larger deep
coral reef formations have been successfully
identified from multibeam imagery (Roberts et
al. 2005), the low resolution of surface-mounted
sonar will hinder efforts to identify some coral
habitats using this technology alone.

The Gulf of Mexico, Pacific coast, and Alaskan
regions have the most extensive multibeam
mapping information. Much of the mapping in
the Gulf of Mexico and the West Coast regions
was conducted as part of oil and gas exploration
activities, while mapping in Alaska has been

undertaken primarily in association with biological
studies or for navigational purposes. Recently,
some deeper water areas around the Main
and Northwestern Hawaiian Islands, American
Samoa, and other U.S. Pacific territories have
been mapped (Chapter 4; Miller et al. 2003) by
NOAA. Likewise, important mapping efforts are
underway in the Gulf of Maine on the northeast
U.S. shelf. In this region, anticipated multibeam
mapping of the continental slope and canyons
will reveal bottom topography and substrates
most likely to support corals, thus allowing
more efficient and directed sampling efforts. A
comprehensive effort to use existing habitat maps
to predict the location of deep coral habitats has
not yet occurred in any region.

As noted above, the South Atlantic Bight has the
most extensive deep coral reefs known to date in
U.S. waters. However, with the exception of the
relatively shallow Oculina banks (Figure 1.31),
there is no synoptic multibeam bathymetry and
backscatter imagery for the shelf break, slope,
and Blake Plateau. Given the unique character
of these deep reef habitats and the potential for
identifying coral bioherms, this region is among
the priorities for mapping. Limited mapping in this
region was conducted in 2007. Since National
Marine Sanctuaries also have the authority
and responsibility to preserve deep coral
communities within their boundaries, mapping,
and characterizing deep coral communities in
the sanctuaries is a priority.

In addition to broad-scale habitat mapping efforts,

Chapman's Reef: Oculina HAPC

&n-4rl MW UNCW

Figure 1.31 3-D colored bathymetry of Chapman's Reef, from 2005 survey done with multibeam
sonar from RN Cape Fear by Seafloor Systems, Inc. Image credit: A. Maness.


focused fine-scale mapping of known deep coral
areas is needed, using side-scan sonar and
in situ ground-truthing (e.g., submersibles or
ROVs). State-of-the-art technologies, such as
autonomous underwater vehicles (AUVs) and
laser-line scan also show promise for finer scale
mapping and habitat characterization.

Modeling the Distribution of
Deep Coral Habitats

Even with detailed multibeam maps of the
seafloor, researchers, and managers will still
be severely limited by the high costs of ground-
truthing potential deep coral areas. Therefore,
alternative techniques for targeting finer-scale
studies will be needed. One promising approach
involves modeling coral habitat requirements
coupled with validation from in situ observations.
Factors to be modeled may include substrate
type, seafloor geomorphology, hydrography,
nutrient levels, and water temperature (Freiwald
et al. 2004). For example, Leverette and Metaxas
(2005) used predictive models to identify suitable
habitat for Paragorgia arborea and Primnoa
resedaeformis, two major structure-forming
gorgonians in the Canadian Atlantic continental
shelf and slope. Modeling the distribution of deep
coral habitats will greatly facilitate focusing future
research efforts geographically and to identify
areas where a precautionary management
approach is warranted until ground-truthed data
can be collected. The accuracy and efficacy of
such models is dependent on the quality of data
inputs and consequently this approach is still
dependent, to some degree, on costly collection

Data Mining and Data Management

Identification of new deep coral areas will
continue to depend upon visual ground-truthing
in addition to acoustic mapping and modeling.
Because of the cost of new exploratory surveys,
there is a high priority to "mine" data from
museum collections or past submersible surveys
focused on other subjects (e.g., geology or fish)
to yield distributional data for corals at a low
cost. Some of these (e.g., video transects) may
also provide qualitative baselines for assessing
change. NMFS has been conducting trawl
surveys since its inception in the 1970's and
much could be learned from this existing data
source. A new Southeastern Area Deep Sea

Coral initiative has begun to systematically
document the distribution of deep corals in the
South Atlantic Bight based on existing data
collected during NOAA-sponsored submersible
and ROV operations.

There is also a need to better manage existing
information to enhance research collaboration
and access to data for management purposes.
The South Atlantic Fishery Management Council,
in coordination with the Florida Wildlife Research
Institute and NOAA, has experimented with web-
accessible data models to combine deep coral
data and other ecosystem information for the
Southeast U.S. region. NOAA is collaborating
with the U.S. Geological Survey and the United
Nations Environment Programme's World
Conservation Monitoring Centre in new deep
coral database efforts. NOAA's Coral Reef
Information System (CoRIS), primarily dedicated
to serving shallow-water coral reef data and
information, currently contains deep coral
information submitted on an ad hoc basis, but
has indicated its interest in expanding efforts to
serve deep coral data.


Monitoring is key to understanding the state
of resources and gaining clues to processes
that may effect change. The United States
identified the development and implementation
of a nationally coordinated, long-term program
to monitor shallow-water tropical reefs as a
key conservation objective (USCRTF 2000). In
contrast to shallow reefs, where a national coral
reef monitoring network is taking shape (Waddell
2005), the costs associated with assessment
and monitoring in the deep sea are much higher.
As a result, it is likely that many deep coral
communities remain to be discovered, baseline
data are limited for most known occurrences,
and quantitative repeated measures are largely

To date, monitoring of deeo croals in U.S. waters
has been limited to select locations off Hawaii
and the southeast U.S. In Hawaii, monitoring
has concentrated on species targeted for harvest
(primarily black and pink corals), but has yielded
valuable life history and ecological information
on these corals (Chapter 4). An infestation of
the invasive snowflake coral, Carijoa riisei, was
also incidentally discovered during monitoring


efforts and is now a major factor shaping recent
management and harvest decisions. Systematic
monitoring of the Oculina Banks Experimental
Research Reserve, a 315 km2 subset of the 1,043
km2 Oculina Bank HAPC, was initiated in 2005.
Between 1994 (when all fishing for snapper and
grouper species was prohibited in the Reserve)
and 2004, 56 ROV dives and 15 research
submersible dives had explored only 0.11%
of the HAPC. In 2005, regular observations on
baseline transects at the same sites in protected
and recently discovered unprotected banks were
initiated (M. Miller pers. comm.). Although it is
too early to assess the success of this approach,
this appears to be the first effort to systematically
monitor a deep coral reef ecosystem. The South
Atlantic Fishery Management Council developed
an Oculina Research and Evaluation Plan (http://
ocean.floridamarine.org), but funding for follow-
on monitoring has not been identified.

Taxonomy, Biology, and Life History of
Deep Coral Species

Despite recent advances in the study of deep
coral taxa, much of their basic life history and
biology is still unknown. Worldwide, the greatest
emphasis has been placed on studying the
few species of stony corals, such as Lophelia
pertusa, that form deep reef-like structures. In
U.S. waters outside the Southeast and Gulf
of Mexico, the most abundant and important
structure-forming corals are the gorgonians, with
hydrocorals, black corals, and pennatulaceans
providing significant habitat complexity in certain
regions. The basic taxonomy of these deep
coral taxa, their biogeography, and processes
that may contribute to distributions and
endemism are poorly known. Genetic studies of
key structure-forming species can contribute to
understanding both taxonomic relationships and
connectivity among populations. The latter can
provide information to determine larval source-
sink patterns and gene flow between deep
coral populations and is key to understanding
recruitment dynamics.

Basic life history and ecological studies are
needed to contribute to understanding the
population biology, changes in abundance over
time, and factors affecting the resilience of deep

processes of growth and mortality for key coral

Biodiversity and Ecology of Deep Coral

Structure-forming deep corals have been shown
to provide important ecosystem functions in the
deep-sea environment especially as habitat
for numerous other species. With the exception
of Oculina reefs off Florida, the biodiversity
of these communities in U.S. waters has not
been quantitatively assessed, and functional
relationships between the corals and associated
species are incompletely understood. In addition
to species inventories and quantifying the
associations between corals, other invertebrates,
and fish, studies are needed to characterize
trophic dynamics within deep coral communities
and the life history of associated species.

Understanding the ecological function of these
communities, including their role in mediating
patterns of biodiversity and their importance
as habitat for federally managed species,
is a management priority. Designation and
subsequent protection of HAPCs in the United
States depends on a demonstrated linkage
between a federally managed fish species
and deep corals or other associated habitat
features i.e., demonstration that these features
represent EFH as defined by the Magnuson-
Stevens Act. When the Act was reauthorized in
2006, Councils received additional discretionary
authority to designate zones other than EFH for
the protection of deep-sea corals. Under the
National Marine Sanctuaries Act, deep corals can
be preserved for their intrinsic value as sensitive
and important components of the ecosystems
within the sanctuaries.

Effects of Climate Change and Ocean

Deep corals may provide windows into past
environmental conditions in the deep ocean, as
well as clues for prospective analyses of future
changes that may result from climate change.
A growing number of researchers are looking
at isotopic proxies for past temperature or other
environmental conditions over decades in long-
lived gorgonians and over geologic timescales in
stony coral reef mounds (Smith et al. 1999; Risk


Deep coral communities are vulnerable to
changes in ocean chemistry associated with
increased atmospheric CO2 from the combustion
of fossil fuels (Guinotte et al. 2006). There
have been no studies on the sensitivity of deep
corals to CO2-associated ocean acidification,
but potentially calcification rates, especially of
stony corals such as Lophelia will decrease,
and conditions in vast areas of the ocean may
become unsuitable for deep reef accretion (Royal
Society 2005).

Fishery Impacts

From a management perspective, filling
information gaps on human activities that may
impact deep coral communities is a critical need.
Because fishing impacts are currently the major
threat to these communities in U.S. waters
and around the world, it is especially important
to gain a comprehensive understanding of
fishing effort and distribution. Coral bycatch in
fisheries and stock assessments have proven
especially valuable in mapping coral resources
and interactions with fisheries in Alaska and the
West Coast (Chapters 2 and 3). NOAA's long-
standing trawl surveys and observer programs
in the Northeast are well positioned to include
these types of observations and analyses. The
Southeast Region, in both the southeast U.S.
and the Gulf of Mexico, currently needs improved
reporting and mapping of fishing effort, as well
as increased observer coverage, reporting, and
analysis of coral bycatch.

Other Anthropogenic Stressors

A number of other localized anthropogenic
impacts, such as those associated with oil and
gas exploration and development and with cable
and pipeline deployment, have been reported in
deep coral habitats within U.S. waters. Because
the extent and impacts of these stressors
to deep coral communities is incompletely
documented, there is a need is to characterize
the spatial distribution of these impacts and their
ecological consequences. Once this information
is well understood, management plans may
be implemented to relocate these activities to
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Appendix 1.1. This table represents a compilation of the major structure-forming deep coral species found
within the U.S. EEZ in one or more of the Pacific regions. The species were identified by regional authors
based on one or more criteria including abundance, size (>15 cm), and associations with other invertebrates.
* Corals identified by regional authors as major structure-forming species, o Coral species occurring in region
but not identified by regional author as major structure-forming. Coral genus with a species not identified or
not specified may represent different species in the genus in a different region


-ur----------- -u----- ----- ------------



. The species were identified by regional authors
(>15 cm), and associations with other invertebrates.
-forming species, o Coral species occurring in region
rming species. Deep-water corals reported by a
bean only. ~ Indicate strcuture forming coral found in

o O

0 0

0 *
0 O

armatarm t


0 0

Ellisella elongala o *
Vicella obesa *

Riisea paniculata o *
canella arbuscula 0 *

Keratoisis flexibilis o *

Keratoisis spp. o *
Paragorgia arborea *
Swiftia exsedla *
smericana o o 0
americana della

Varella hellissima o *

rVaella pauciflora o *




Robert P. Stone and S. Kalei Shotwell


Alaska is the largest state in the U. S. and contains
more than 70% of the nation's continental shelf
habitat. The state has 55,000 km of tidal shoreline
and the surface area of marine waters in the
U.S. Exclusive Economic Zone (EEZ) measures
approximately 3.3 million km2. The region has a
highly varied submarine bathymetry owing to the
numerous geological and physical processes at
work in the three main physiographic provinces
- continental shelf, continental slope, and
abyssal plain. The marine environment of the
Alaska Region can be divided into three major
geographical subregions the Gulf of Alaska,
the Bering Sea including the Aleutian Island
Archipelago, and the Chukchi and Beaufort Seas
in the Arctic.

Deep corals are widespread throughout Alaska,
including the continental shelf and upper slope
of the Gulf of Alaska, the Aleutian Islands,
the eastern Bering Sea, and extending as far
north as the Beaufort Sea. Coral distribution,
abundance and species assemblages differ
among geographic regions. Gorgonians and
black corals are most common in the Gulf of
Alaska while gorgonians and stylasterids are
the most common corals in the Aleutian Islands.
True soft corals are common on Bering Sea shelf
habitats. Overall, the Aleutian Islands have the
highest diversity of deep corals in Alaska, and
possibly in the North Pacific Ocean, including
representatives of six major taxonomic groups
and at least 50 species or subspecies of deep
corals that may be endemic to that region. In the
Aleutian Islands, corals form high density "coral
gardens" that are similar in structural complexity
to shallow tropical reefs and are characterized

Auke Bay Laboratory, Alaska Fisheries Science
National Marine Fisheries Service
11305 Glacier Highway
Juneau, Alaska 99801-8626

by a rigid framework, high topographic relief and
high taxonomic diversity (Stone 2006).

A few coral species were described from Alaskan
waters as early as the late1800's (Verrill 1865;
Dall 1884), but the true magnitude of Alaska's
coral resources was not realized until the U.S.
Fisheries Steamship Albatross brought back
evidence of rich beds of corals in 1888. The
Albatross Expedition continued through 1906
in Alaskan waters and collections made during
that period initiated the first detailed taxonomic
work on Alaskan octocorals (Nutting 1912) and
hydrocorals (Fisher 1938). With specific regard
to hydrocorals Fisher (1938) noted that "the
North Pacific is far richer in indigenous species
than the North Atlantic." Collections made since
that time, mostly opportunistic rather than from
directed expeditions, have resulted in subsequent
taxonomic work on octocorals (Bayer 1952; Bayer
1982; Bayer 1996), antipatharians (Opresko
2005), and a synthesis on scleractinian corals
(Cairns 1994).

Most information on coral distribution in Alaska is
based on fisheries by-catch and stock assessment
survey data. Consequently, our knowledge of
coral distribution is largely limited, and somewhat
biased, to those geographic areas and depth
zones where fisheries and stock assessment
surveys have occurred. Nonetheless, given
the widespread nature of existing fisheries and
surveys in the state, the distribution of coral
from these sources provides a fairly accurate
depiction of the true distribution of corals. Few
directed studies have been undertaken until
recently to examine the ecology and distribution
of deep corals. Cimberg et al. (1981) compiled
a synthesis of coral records from Alaskan waters
specifically to address concerns about oil and
gas exploration and development on the outer
continental shelf. Some information on coral
distribution has been opportunistically collected
during nearshore scuba and submersible surveys
focused on fish stock assessments, fish habitat



Figure 2.1. Map of Alaska showing the 5 broad geographical areas that were delineated for this report. From
east to west eastern Gulf of Alaska (red box), western Gulf of Alaska (black box), eastern Aleutian Islands
(green box), western Aleutian Islands (purple box), and Bering Sea (blue box).

assessments, and studies on the effects of fishing
gear on fish habitat.

Two major research programs were recently
initiated in largely unexplored areas of Alaska and
findings from those studies, although preliminary,
have greatly increased our knowledge on
the distribution of deep corals. Following an
exploratory cruise in 2002, a multi-year study was
initiated to investigate coral habitat in the central
Aleutian Islands using the manned submersible
Delta and the remotely operated vehicle (ROV)
Jason I/. The National Oceanic and Atmospheric
Administration's National Marine Fisheries Service
(NOAA/NMFS), the North Pacific Research
Board (NPRB), and NOAA's Undersea Research
Program (NURP) sponsored this research. In
2002 and 2004, a multi-discipline study using

the manned submersible Alvin was launched
to investigate seafloor habitat on North Pacific
Ocean seamounts. A total of seven seamounts
within the U. S. EEZ were explored during the
two-year study. An additional seamount located
south of the Alaska Peninsula was explored with
the ROV Jason II in 2004. NOAA's Office of
Ocean Exploration (OE) and NURP sponsored
the seamount studies

In this chapter, detailed descriptions of deep coral
habitat found in Alaskan waters are provided
along with a discussion of their distribution,
threats to deep coral habitat, and current
management and conservation measures. Five
broad geographical areas of Alaska (Figure 2.1)
were delineated as follows: 1) the eastern Gulf of
Alaska (GOA) including the inside waters of the


Alexander Archipelago, Southeast Alaska, 2) the and methane ,
western GOA including the Alaska Peninsula, 3) of the seabed
the eastern Aleutian Islands (Shumagin Islands been strongly
to Seguam Pass), 4) the western Aleutian Islands rates of sedime
(Seguam Pass to Stalemate Bank), and 5) the also contains a
Bering Sea. arranged in th
the Juan de F
Coral records from these areas were categorized volcanoes risin
into the six major taxonomic groups. Three likely formed c
ecologically important groups of gorgonians, mantle hotspot
Primnoa spp., Paragorgia spp., and bamboo
corals(Familylsididae) arecategorizedseparately The Bering Sei
because their large size and conspicuous The Bering Se.
morphology greatly reduce the probability of the largest cont
inaccurate field identification. km long and 5(
Sciences 199C
The principal source of information on coral at approximate
distribution is by-catch data collected during NMFS canyons, inclu
research trawl surveys (Resource Assessment Canyons-the

(RACEBASE)), Alaska Fisheries Science
Center (AFSC), Resource Assessment and
Conservation Engineering Division's Groundfish
Assessment Program). Although RACEBASE
includes records of research cruises since 1954,
data collected prior to 1975 are not included in
this report because the catch of corals was not
always recorded and the accuracy of onboard
coral identifications made before that time is
questionable. By-catch data collected during the
AFSC sablefish longline survey in 2004, published
records, and unpublished in situ observations
were also used to map coral distributions. There
is very limited survey and fishery information from
the Alaskan Arctic (Chukchi and Beaufort Seas).


The Gulf of Alaska
The Gulf of Alaska has a broad continental shelf
extending seaward up to 200 km in some areas
and contains several deep troughs (National
Academy of Sciences 1990). In the eastern
Gulf of Alaska, the Pacific Plate moves roughly
parallel to the North American Plate, along the
Fairweather-Queen Charlotte fault, and forms an
abrupt continental slope with an abbreviated shelf
(NURP 1996). In the northern and western parts
of the Gulf of Alaska, the two plates slide, rather
than slip past each other, and form a convergent
margin and subduction zone (NURP 1996). Gulf
of Alaska continental shelf habitats include steep
rock outcrops, smooth turbidite sediment scapes,

)eps (NURP 2001). The nature
an the Gulf of Alaska shelf has
ifluenced by glaciation and high
it deposition. The Gulf of Alaska
proximately 24 major seamounts
ee chains extending north from
ca Ridge. The seamounts are
from the abyssal plain that were
s the Pacific Plate moved over

is a shallow sea and has one of
mental shelves in the world 1200
3 km wide (National Academy of
. The continental shelf breaks
/ 170 m depth and seven major
ing the Zhemchug and Bering
wo largest submarine canyons
(Normark and Carlson 2003),

indent the continental slope (Johnson 2003).
The continental shelf is covered with sediment
deposited by the region's major rivers (Johnson
2003) and therefore has limited hard substrate for
coral attachment. TheAleutian IslandArchipelago
contains more than 300 islands and extends
over 1900 km from the Alaska Peninsula to the
Kamchatka Peninsula in Russia. TheArchipelago
is supported by the Aleutian Ridge and it forms a
semi-porous boundary between the deep North
Pacific Ocean to the south and the shallower
Bering Sea to the north. The Aleutian Ridge is a
volcanic arc with more than 20 active volcanoes
and frequent earthquake activity that was formed
along zones of convergence between the North
American Plate and other oceanic plates (Vallier
et al. 1994). The island arc shelf is very narrow
in the Aleutian Islands and drops precipitously on
the Pacific side, to depths greater than 6000 m in
some areas, such as the Aleutian Trench.

The Alaskan Arctic
The Bering Strait separates the Bering Sea from
the Chukchi Sea. The Chukchi Sea is a shallow
shelf (only 20 to 60 m deep). The continental shelf
in the Beaufort Sea is fairly broad (80-140 km
wide) and is a submarine extension of the North
Slope coastal plain (Horowitz 2002). Sediments
on the continental shelf are predominantly soft
and fine-grained and are redistributed by long-
shore currents, wave action, entrainment in
bottom-fast ice, ice gouging, ocean currents, and
internal waves (Horowitz 2002).



Major oceanic currents are found in all three
subregions of Alaska and variations in their
circulation control the climate and oceanic patterns
in the North Pacific and Arctic Oceans. Currents
likely influence larval dispersal and consequently
the distribution of deep corals. Major oceanic
currents influence the water temperature regimes
in the subregions that may affect the growth rates
for some species of deep corals.

The Gulf of Alaska
Two primary ocean currents exist in the Gulf of
Alaska that flow around the Alaska Gyre. The
Alaska Current is a wide (>100 km), slow moving
(0.3 m s-1) current that flows northward off the
shelf of the eastern Gulf of Alaska. It becomes
the Alaska Stream west of Kodiak Island where it
narrows (<60 km), increases speed (1 m s-1) and
continues to flow westward south of the Alaska
Peninsula and Aleutian Island Archipelago (Royer
1981). Continental shelf circulation is strongly
influenced by freshwater input, and nearshore
currents are additionally influenced by shelf
bathymetry (Allen et al. 1983). Some areas of
the Gulf of Alaska have among the largest tides
in the world (Cook Inlet has the 2nd largest tidal
amplitude in NorthAmerica, after the Bay of Fundy
in Atlantic Canada) and circulation is strongly
tidally influenced in those areas. Several physical
processes enhance regional nutrient supply and
primary productivity and include costal upwelling,
river discharge, tidal mixing, estuarine circulation,
mesoscale eddy formation and transport, and
recirculation around the Alaska Gyre (Whitney et
al. 2005).

The Bering Sea
The Aleutian Archipelago forms the boundary
between the deep North Pacific Ocean and
the shallower Bering Sea. Deep water flowing
northward in the Pacific Ocean encounters the
Aleutian Trench where it is forced up onto the
Aleutian Ridge and into the Bering Sea through
the many island passes (Johnson 2003).
Additionally, coastal water from the Alaska
Stream enters through Unimak Pass in the
eastern Aleutians and slowly (0.01 to 0.06 m s-1)
flows northeastward along the Alaska Peninsula.
The Aleutian North Slope Current flows eastward
on the north side of the Aleutian Islands towards
the inner continental shelf of the Bering Sea.
This is a swift current (0.5 m s-1) and the steep

continental slope forces much of the flow into the
northwest flowing Bering Slope Current (Johnson

The Bering Slope Current flows northwestward
off the shelf break and together with currents on
the northern shelf flows northward through the
Bering Strait and into the Chukchi Sea (Kinder
and Schumacher 1981). Tidal currents dominate
circulation in the southeastern shelf area of the
Bering Sea (Kinder and Schumacher 1981). On
the outer shelf currents flow along isobaths to the
northwest at speeds up to 0.1 m s-1.

The Alaskan Arctic
North Pacific waters flow from the Bering Sea,
across the Bering Strait and into the Chukchi Sea
in the Arctic. Consequently, the Chukchi Sea has
more faunal affinities to the North Pacific than to
the deeper Beaufort Sea. Different circulation
regimes exist on the inner and outer continental
shelves of the Beaufort Sea (Aagaard 1984).
Circulation on the inner shelf is to the west and
strongly wind-driven. Outside the 50-m isobath,
the Beaufort Undercurrent slowly (0.1 m s-1) flows


Coral communities in Alaskan waters are highly
diverse and include six major taxonomic groups
(Appendix 2.1): true or stony corals (Order
Scleractinia), black corals (Order Antipatharia),
true soft corals (Order Alcyonacea) including
the stoloniferans (Suborder Stolonifera), sea
fans (Order Gorgonacea), sea pens (Order
Pennatulacea), and stylasterids (Order
Anthoathecatae). One hundred and forty one
unique coral taxa have been documented from
Alaskan waters and include 11 species of stony
corals, 14 species of black corals, 15 species of true

soft corals (including six species of stoloniferans),
63 species of gorgonians, 10 species of sea
pens, and 28 species of stylasterids (Appendix
2.1). Note that all taxa listed in Appendix 2.1 are
believed to be unique and include 52 taxa with
incomplete taxonomy, including several that have
only recently been collected and likely represent
species new to science. All corals found in
Alaska are azooxanthellate and satisfy all their
nutritional requirements by the direct intake of
food. They are ahermatypic or non-reef building


corals but many are structure forming. The
degree to which they provide structure depends
on their maximum size, growth form, intraspecific
fine-scale distribution, and interaction with other
structure-forming invertebrates (Table 2.1).

a. Stony corals (Class Anthozoa, Order
At least 11 species of stony corals have been
reported from Alaskan waters (Cairns 1994).
All are solitary cups and the largest species
measure less than 10 cm in total height. They
require exposed, hard substratum for attachment.
Unlike their tropical counterparts, they do not
form significant structure used by larger fishes
as refuge (Table 2.1). They are, however,
contagiously distributed (i.e. aggregated or
clumped) and dense patches may provide some
structural habitat for some macro-invertebrates
and juvenile fish (Figure 2.2).

b. Black Corals (Class Anthozoa,
Order Antipatharia)
Black corals have some importance as structure-
forming corals (Table 2.1) and at least 14 species

Table 2.1. Structure-forming attributes of deep corals in

Figure 2.2. Scleractinians occasionally form
dense patches, such as this one in Amchitka
Pass (Aleutian Islands), that may provide refuge
habitat for small fish and crustaceans. Photo by
R. Stone, NOAA Fisheries.

are reported from Alaska (Appendix 2.1). They
are locally abundant, contagiously distributed,
and a few species such as Dendrobathypathes
boutillieri (Opresko 2005) and Parantipathes
sp. may grow over 1 m in height and/or width
(Figure 2.3). Data from the NMFS sablefish
longline survey indicate that several species

No Low Small No-branch Few Clumped Low
No Medium Large Branch Few Clumped Medium
No Medium Small No-branch Few Clumped Medium
No Low Small No-branch Few Clumped Low
No High Large Branch Many Clumped High
No High Large Branch Many Clumped High

No Medium Large Branch Many Clumped High
No Medium Large Branch Many Clumped High
No High Large Branch Few Clumped Medium
No High Medium Branch Many Clumped High

Small (<30cm)/ Medium (30cm-1m)/ Large (>1m)
Branching/ Non-branching
None/ Few (1-2)/ Many (>2)


Figure 2.3. Some black corals such as this Dendro-
bathypathes boutillieri may reach heights over 1 m.
An unknown species of octopus takes cover under the
coral. Photo credit: R. Stone, NOAA Fisheries.

form dense patches in some areas of the Gulf of
Alaska. Deep ROV observations in the central
Aleutian Islands in 2004 confirmed that black
corals are contagiously distributed with densities
approaching 1 colony m-2 on some shelf habitats
(R. Stone, unpublished data). They require
hard substratum for attachment and by-catch
specimens collected during NMFS groundfish
surveys in the Gulf of Alaska were attached to
small cobbles and mudstone.

c. Gold Corals (Class Anthozoa, Order
Gold corals or zoanthids are not known to occur
in Alaskan waters but dense mats of zoanthid-like
colonies similar to Epizoanthus scotinus known
from British Columbia (Lamb and Hanby 2005)
have been observed in eastern Gulf of Alaska
habitats (R. Stone, personal observations).

d. Gorgonians (Class Anthozoa,
Order Gorgonacea)
Gorgonians are the most diverse coral group
in Alaskan waters more than 60 species
representing seven families have been reported

(Appendix 2.1). Gorgonians are also the most
important structure-forming corals in Alaskan
waters (Table 2.1). They generally require
exposed, hard substratum for attachment but
recent observations in deep water (>450 m)
indicatethatthe skeletons of hexactinellid sponges
may be important attachment substrates in areas
devoid of exposed rock (R. Stone, unpublished
data). Gorgonians are locally abundant,
contagiously distributed, and several species
attain massive size. Gorgonians form both single-
and multi-species assemblages. For example,
Primnoa pacifica forms dense thickets in the
Gulf of Alaska (Krieger and Wing 2002) while as
many as 10 species are found in Aleutian Island
coral gardens (Stone 2006). Some gorgonians
are also extremely long lived. A medium-sized
colony (197.5 cm length) identified as Primnoa
resedaeformis (most likely P pacifica) was aged
at 112 years in the Gulf of Alaska (Andrews et al.
2002). P, pacifica attains a height of 7 m in the
Gulf ofAlaska (Krieger2001) and P wingi reaches
a height of at least 1.5 m in the Aleutian Islands
(R. Stone personal observations). The depth
and geographical distribution of Primnoa spp. in
Alaskan waters corresponds to the mean spring
bottom temperature of 3.70C (Cimberg etal. 1981)
suggesting that this might be the low temperature
of its tolerance range. Paragorgia arborea can
measure 2 m high and wide, (Figure 2.5) and other
gorgonians such as Plumarella sp., Fanellia sp.,
and bamboo corals (Family Isididae) grow to over
1 m high (R. Stone personal observations). The
northern distribution of bamboo corals suggests

Figure 2.4. This true soft coral (Anthomastus sp.)
measures 20 cm across and provides shelter for a
snailfish (Careproctus sp.). Photo credit: R. Stone,
NOAA Fisheries.


a temperature tolerance of less than 3.0C and
their distribution also suggests a low tolerance
for high sedimentation (Cimberg et al. 1981).

e. True Soft Corals and Stoloniferans
(Class Anthozoa, Order Alcyonacea)
True soft corals (Suborder Alcyoniina) are not
a diverse group in Alaskan waters only nine
species are reported (Appendix 2.1). They have
some importance as structure-formers (Table
2.1). Colonies are encrusting or erect and a
few species (e.g., Anthomastus ritterii) may
reach 20 cm in height (Figure 2.4) They require
exposed, hard substratum for attachment,
are locally abundant, and have a contagious
distribution. Eunephthea rubiformis (formerly
Gersemia rubiformis) are locally abundant on
the unconsolidated sediments of the eastern
Bering Sea shelf (Heifetz 2002) and although
small, colonies may be abundant enough to
provide important refuge habitat for juvenile fish
and crustaceans. Additionally, six species of
stoloniferans (Suborder Stolonifera) are reported
from Alaska (Appendix 2.1) and they generally
have little importance as structure-formers (Table
2.1). They can form extensive mats on hard
surfaces such as rock, other corals, and sponges
(Stone 2006). They are locally abundant a
single species of Clavularia was measured at a
density of 1.7 colonies m-2 in one Aleutian Island
coral garden (Stone 2006).

f. Pennatulaceans (Class Anthozoa, Order
Ten species of pennatulaceans (sea pens) are
reported from Alaskan waters (Appendix 2.1)
and several are important structure-forming

Figure 2.6. Dense groves of the sea pen Ptilosarcus
gureyi are found on soft-sediment shelf habitats in
the Gulf of Alaska and Aleutian Islands. Photo credit:
P. Malecha, NOAA Fisheries.

Figure 2.5. A large bubblegum coral (Parago-
rgia arborea) provides shelter for a Pacific cod
(Gadus macrocephalus) in the central Aleutian Is-
lands. Photo credit: R. Stone, NOAA Fisheries.

corals (Table 2.1). Many species are elongate
and whip-like and one species, Halipteris
willemoesi, attains a height greater than 3 m (R.
Stone personal observations). At least three
species form extensive groves in soft-sediment
areas. Protoptilum sp. and H. willemoesi form
dense groves (16 m-2 and 6 m-2, respectively)
in the central Gulf of Alaska (Stone et al. 2005).
Dense groves of H. willemoesi have also been
reported on the Bering Sea shelf (Brodeur 2001).
Ptilosarcus gurneyi also forms dense groves
on shallow shelf habitats throughout the Gulf of
Alaska and Aleutian Islands (Figure 2.6).

g. Stylasterids (Class Hydrozoa,
Order Anthoathecatae)
More than 25 species or subspecies are reported
from Alaskan waters (Wing and Barnard 2004;
Appendix 2.1) and many are important structure-
forming corals (Table 2.1). They form erect (e.g.,
Stylaster spp.) or encrusting calcareous colonies
(e.g., Stylantheca petrograpta), and require
exposed, hard substratum for attachment (Figure
2.7). Some erect species, most notably Stylaster
cancellatus, may grow to almost one meter in
height and often display contagious distributions.
Stylasterids, particularly Stylaster campylecus,
are a major structural component of Aleutian
Island coral gardens and are often encrusted with
the demosponge Myxilla incrustans together
they form a rigid platform that other sedentary and
sessile invertebrates use as an elevated feeding
platform (Stone 2006). Encrusting species, such
as S. petrograpta, have low value as structure-
forming invertebrates.



Deep corals are widespread in Alaska and have
been reported as far north as the Beaufort Sea
(Cimberg et al. 1981). Corals are found over a
broad depth range and occur from the shallow
subtidal zone to the deep ocean trenches (Table
2.2). For example, pennatulaceans have been
found as shallow as 3 m depth and antipatharians
and gorgonians have been found at a depth of
4784 m on Gulf of Alaska seamounts. They are
found in all megahabitats and mesohabitats as
described by Greene et al. (1999). In addition to
general factors controlling coral distribution such
as current regimes and the presence of hard
substrates, temperature tolerance appears to play
a role in the geographic and depth distribution of
some deep corals.

Eastern Gulf of Alaska
Deep corals have a widespread but patchy
distribution on the continental shelf and slope
in the eastern Gulf of Alaska (Figure 2.8).
Approximately 46 species are reported from the
area (Appendix 2.1). Only the Aleutian Islands
support a higher diversity of corals. Corals
include four species of stony corals, nine species
of black corals, four species of true soft corals
(including two stoloniferan species), thirteen
species or subspecies of gorgonians, seven
species of pennatulaceans, and nine species or

Figure 2.7. Large, erect stylasterids (Stylastersp).
grow on exposed bedrock with their central axis per-
pendicular to the current in the Aleutian Islands. Red
laser marks are separated by 10 cm. Photo credit:
R. Stone, NOAA Fisheries.

subspecies of stylasterids (Appendix 2.1).

Corals range in depth from 6 m forPrimnoapacifica
in the glacial fiords of Glacier and Holkham Bays
(Stone et al. in preparation) to over 400 m on the
continental slope. P pacifica is found throughout
the subregion and forms dense thickets in
some areas, especially in the inside waters of
Southeast Alaska and on high-relief rocky areas
of the continental shelf (Figure 2.8A). It grows on
bedrock and boulders and has been observed in
situ at a depth of 365 m (Krieger2001). Anecdotal
information exists that it may grow as deep as 772
m in some areas of Southeast Alaska (Cimberg
et al. 1981). Stylasterids are fairly common on
the continental shelf and in some shallow areas
of Southeast Alaska (Figure 2.8B). Black corals
grow on the continental shelf at depths between
401 and 846 m (Figure 2.8C). Stony corals and
soft corals are known from only a few locations
(Figure 2.8D and 2.8E).

Calcigorgia spiculifera is another important
gorgonian in Southeast Alaska that forms small
groves on bedrock in shallow water areas (Stone
and Wing 2001). The pennatulaceans, Halipteris
willemoesi and Ptilosarcus gurneyi also form
dense groves in some areas (Figure 2.8F) at
depths between 20 and 200 m (Malecha et al.
2005). The most ecologically important coral
feature in this subregion of Alaska is the Primnoa
thickets on the continental shelf of the eastern
Gulf of Alaska (Figure 2.8A). In July 2006,
NMFS closed five small areas where Primnoa
thickets have been documented via submersible
observations to all fishing activities using bottom-
contact gear.

Western Gulf of Alaska
Deep corals have a widespread but patchy
distribution in the western Gulf of Alaska (Figure
2.9). Gorgonians are widely distributed on the
continental shelf and slope (Figure 2.9A) and
are represented by 13 species (Appendix 2.1).
Primnoa sp. is the most common gorgonian with
unconfirmed reports of dense thickets in the
area of Chirikof Island (Cimberg et al. 1981).
Bamboo corals are patchily distributed on the
continental slope and records of Paragorgia spp.
are rare (Figure 2.9A). Stylasterids are widely
distributed (Figure 2.9B) but are not abundant or
diverse. Only two species have been reported
from this subregion (Appendix 2.1). Black
corals, stony corals, and soft corals have only


14.j W

130 W

140 W


-- 1000M (q
Xr stow CQTW V
0 100 2 0

o 2o% Joo


140 W 130 W

140'11 1401W

Figure 2.8. Distribution of corals in the eastern Gulf of Alaska A) gorgonians (bamboo corals Family Isidi-
dae, Paragorgia spp., Primnoa spp. are plotted separately), B) stylasterids, C) black corals, D) stony corals, E)
soft corals, and F) pennatulaceans.



I10 a'

130 W


Table 2.2. Summary of species richness and depth range for seven major
groups of corals found in Alaskan waters. Data sources for depth distribution: 1
A. Baco-Taylor, unpublished data; 2. Hoff and Stevens 2005; 3. Keller 1976; 4.
R. Stone, unpublished data; 5. Stone et al. in preparation; 6. Stone 2006.

11 24 4620 4-3
14 401-4784 4-1
9 10-3209 4-2
6 11 -591 6-4
63 6 4784 5- 1
10 3 2947 4-4
28 11 -2130 6-4
141 3 4784

been reported from a few areas (Figures 2.9C,
2.9D, 2.9E). The most ecologically important
coral feature in this subregion of Alaska is the
extensive pennatulacean groves (Figure 2.9F) in
the submarine gullies south and east of Kodiak
Island (Stone et al. 2005) and in isolated locations
in Prince William Sound (Malecha et al. 2005).

Gulf of Alaska Seamounts
Submersible observations in 2002 and 2004
confirmed by-catch records that seamounts in the
Gulf of Alaska are rich in coral habitat and that all
major taxonomic groups except stylasterids were
present (Appendix 2.1) (A. Baco-Taylor, WHOI,
pers. comm.). The absence of stylasterids from
the Gulf ofAlaska seamounts is notable since they
are common on the seamounts near New Zealand
(Cairns 1991; Cairns 1992). Pennatulaceans are
also noticeably uncommon from the seamounts
and are represented by a single unidentified
species (Appendix 2.1). The submersible Alvin
was used during a 2004 research cruise to five
seamounts in the northern Gulf ofAlaska (Dickens,
Denson, Welker, Giacomini, and Pratt) to collect
video footage and specimens on transects along
three depth strata: 700 m, 1700 m, and 2700 m.
Corals were most abundant near the seamount
summits (700 m) where Paragorgia spp. and
bamboo corals were the dominant coral fauna.
Gorgonians (Primnoidae) were the mostabundant
corals at the 2700 m depth stratum. Corals were
least abundant and diverse in the 1700 m depth
zone where black corals and Primnoidae were
dominant. Precious red coral (Corallium sp.) was
collected from Patton Seamount and represented

a northern range extension
for the family Corallidae.
Bamboo corals were
a particularly diverse
group with at least four
genera collected on the
seamounts (P. Etnoyer,
Texas A&M University
- Corpus Christi, pers.

Coral habitat on Derickson
Seamount which crests at
2766 m south of theAlaska
Peninsula was explored
with the ROV Jason II
in 2004. Black corals,
bamboo corals, and other
gorgonians (Primnoidae

and Chrysogorgiidae) were observed on hard
substrates at depths between 2766 and 4784 m
(A. Baco-Taylor, WHOI, pers. comm.). Several
specimens collected on this deep seamount
represent species new to science and significant
depth-range extensions. A single species of
stony coral (Fungiacyathus sp.) was observed
in soft-sediment areas. Species distribution
differed between the eastern and northern flanks
of the seamount and highlights the importance
of circumnavigating seamounts during surveys of
coral distribution.

The Aleutian Islands
The Aleutian Islands support the most abundant
and diverse coral assemblages in Alaska
(Appendix 2.1). A total of 101 coral species or
subspecies have been reported from the Aleutian
Islands (Appendix 2.1). Previous reports
indicated that 25 coral taxa were endemic to
the region (Heifetz et al. 2005) our updated
records however, indicate that as many as 51
species may be endemic to the region! Deep-
water collections made with the ROV Jason II in
2004 may add dozens of corals novel species
and range extensions to this list. Gorgonians
and stylasterids are the most diverse groups
with 45 and 25 species or subspecies reported,
respectively (Appendix 2.1). Twelve species
of true soft corals including three species of
stoloniferans, six species of pennatulaceans,
and ten species of stony corals have also
been reported from the subregion (Appendix
2.1). Additionally, three species of black corals
were collected from the area in 2004 (R. Stone,


160 W 150 W

60N n-

Bashn mxI
-- n

S. Island
55 N S 53-N
S-- Isand O isa

150 "A

160 W 150 W

-- I1rllin

X -Slon Co

S55 N -55'N
mzIs land O

160 \\ 150 W

160W 150 W

"lll nim

Illll in i
.0 IslandI
ora 55 N e55 N

Figure 2.9. Distribution of corals in the western Gulf of Alaska A) gorgonians (bamboo corals Family Isidi-
dae, Paragorgia spp., Primnoa spp. are plotted separately), B) stylasterids, C) black corals, D) stony corals, E)
soft corals, and F) pennatulaceans.


unpublished data) including Dendrobathypathes
boutillieri, a species new to science (Opresko

Eastern Aleutian Islands
Data from NMFS stock assessment surveys
indicate that a major shift in coral diversity occurs
in the eastern Aleutian Islands at about longitude
1690 Wnearthe west end of Umnak Island (Heifetz
et al. 2005). Approximately twelve species of
stylasterids, nine species of gorgonians, and
three species of stony corals found further west
in the Aleutian Islands are not found east of this
area (Heifetz et al. 2005).

Gorgonians are widely distributed on the
continental shelf and upper slope (Figure
2.10A). Primnoa spp. and Paragorgia spp. are
widely distributed but few bamboo corals have
been reported from the area (Figure 2.10A).
Stylasterids are widely distributed especially
along the south side of the archipelago (Figure
2.10B). Few black corals have been reported
(Figure 2.10C) but stony corals and soft corals
are widespread and abundant in some areas
(Figures 2.10D, 2.10E). Pennatulaceans are
widely distributed and likely form dense groves in
some areas (Figure 2.10F).

Western Aleutian Islands
Corals are abundant and widespread in the
western Aleutian Islands (Figure 2.11). Coral
gardens, a previously undocumented habitat
feature in the North Pacific Ocean, were observed
with the submersible Delta at six locations in
the central Aleutian Islands during 2002 (Stone
2006). Gardens are typically located in small,
discrete patches at depths between 117 and 338
m and are distinguishable from other habitats by
extremely high coral abundance (3.85 corals
m-2), especially gorgonians (1.78 colonies m-2),
and stylasterids (1.46 colonies m-2).

In general, corals appear to have a much
broader depth distribution in the western Aleutian
Islands than elsewhere in Alaska. The depth
distribution of Primnoa spp. (304-1436 m) is
substantially deeper than elsewhere in Alaska
(Stone 2006; R. Stone, unpublished data).
Bamboo corals and Paragorgia spp. also have
a very broad geographical distribution (Figure
2.11A). Bamboo corals have been observed
at depths between approximately 400 and 2827
m (R. Stone, unpublished data) and have been

collected with a beam trawl at a depth of 3532
m (Cimberg et al. 1981). Paragorgia spp. has
been observed in situ at depths between 27 m
(Stone 2006) and 1464 m (R. Stone, unpublished
data). Stylasterids are widespread (Figure 2.11B)
and have been observed at depths between 11 m
(Stone 2006) and 2130 m (R. Stone, unpublished
data). Black corals appear to have a limited
distribution (Figure 2.11C) and have been
observed on bedrock, boulders, and cobbles
at depths between 449 and 2827 m (R. Stone,
unpublished data).

Stony corals have a fairly broad distribution in
this region of Alaska (Figure 2.11D) and have
been collected at depths between 24 m (R.
Stone, unpublished data) and 4620 m in the
Aleutian Trench (Keller 1976). True soft corals
are also fairly common in this region of Alaska
(Figure 2.11E) and have been observed at
depths between 10 m and 2040 m (R. Stone,
unpublished data). Pennatulaceans have been
observed as deep as 2947 m and form extensive
groves in some soft-sediment areas on both shelf
and slope habitats (Figure 2.11F).

The Bering Sea
Deep corals have a patchy distribution in this
region of Alaska and are largely limited to the
broad, shallow continental shelf and along the
narrow continental slope (Figure 2.12). The
entire north side of the Aleutian Archipelago
is technically within the Bering Sea but for the
purposes of this reportwe have defined the Bering
Sea as those areas of the shelf and slope not
immediately adjacent to the Aleutian Islands (as
illustrated in Figure 2.12 and including the inner
shelf illustrated in Figure 2.9). This definition
applies both to the discussions in the text and to
the species list provided in Appendix 2.1.

The coral fauna of this region of Alaska has
been poorly documented but does not appear
to be particularly diverse. Sixteen species or
subspecies of coral are known from the region
and include three species of true soft corals
(including one species of stoloniferan), six species
of gorgonians, four species of pennatulaceans,
and three species of stylasterids (Appendix
2.1). Additionally, at least one species of black
coral, one species of stony coral, and one
species of bamboo coral have been collected
from the region but proper identifications were
never made. These records effectively increase


1701W 160 W

Unmak Island 5J5,-^ '* N_.

Ba.memlr + Bamboo Conl
-- 200m I I'araonia
1000 m 8 Prinoa
oN- Grcgonians
0 2Wm EEZ
17 W 160 W

1", I0 W

1000 m

170-W 160 W

1701%1" 160 'W
Umnak Island


200m O

o o n.o EEZ
170-W 160 W

Figure 2.10. Distribution of corals in the eastern Aleu
dae, Paragorgia spp., Primnoa spp. are plotted separ,
soft corals, and F) pennatulaceans.
soft corals, and F) pennatulaceans.

170VW 160 W

--1000 m
5ON- 0 Hvdrocoral
o I 2o0 km EEEZ
170 W 160 W

717 0W 160 W

U IslandBt _

1000 m
5oN- 3 Stony Coral

170 W 160'W

1701W 160'W
:F % :
Unmik Island 5Y_ i" vr 1 'SS^N

-- 1000 m
'N- SPennoaluacean
0o Loo San EEZ

170 W 160 W

i Islands A) gorgonians (bamboo corals Family Isidi-

y), B) stylasterids, C) black corals, D) stony corals, E)
y), B) stylasterids, C) black corals, D) stony corals, E)


180 180

Bowees Ridge Boweos Ridge

200 m Bathme rv
1000m 200m
N B mboo Coral 1000m
A Parargia rII Hydrocoral

o0 10 2o00 o 10 200
180 180

I 8: 180

Bowers Ridge Bowers Ridge
su5cN y uN

an ma pr Im

200m 200 m
50N SO 50 N
lo- O Om om 1000m
A Black Coral I Stony Coral

0 l 00 100 0 100 ; 3
180' 1t80

180 180

Bowers Ridge Bowers Ridge

200 mi 200:
1000 m 1000 M
d e n So bCoral Pematioaceans

0 I'l 200 0 I1) 20 I

Figure 2.11. Distribution of corals in the western Aleutian Islands A) gorgonians (bamboo corals Family Isidi-
dae, Paragorgia spp., Primnoa spp. are plotted separately), B) stylasterids, C) black corals, D) stony corals, E)
soft corals, and F) pennatulaceans.


Figure 2.12. Distribution of corals in the Bering Sea A) gorgonians (bamboo corals Family Isididae, Parago-
rgia spp., Primnoa spp. are plotted separately), B) stylasterids, C) black corals, D) stony corals, E) soft corals,
and F) pennatulaceans.


the number of species in the region to at least
nineteen. Gorgonians are distributed mostly on
the continental slope and a few isolated shelf
locations (Figure 2.12A). Primnoa pacifica,
bamboo corals, and Paragorgia sp. have been
collected from a few locations on the continental
slope (Figure 2.12A). The bamboo coral
specimens were collected during NMFS surveys
and because definitive species identifications
were not made they are not included in the
species list (Appendix 2.1). Stylasterids have
been reported from only a single location in the
Pribilof Islands area (Figure 2.12B). Black corals
have been reported from only a single location
on the outer continental slope (Figure 2.12C) and
stony corals are known from a few locations on
shelf and slope locations (Figure 2.12D). The
pennatulacean H. willemoesi forms dense groves
on the outer continental shelf of the Bering Sea
(Figure 2.12F) at depths between 185 and 240
m (Brodeur 2001; Malecha et al. 2005). The
most important coral feature of the Bering Sea
however, is likely the dense aggregations of
soft corals (mostly Eunephthea rubiformis) on
the unconsolidated sediments of the continental
shelf (Figure 2.12E).

Alaskan Arctic
Only the soft coral Eunephthea sp. has been
reported north of the Bering Sea (Cimberg et al.
1981). Eunephthea sp. is patchily distributed on
the shallow shelves of the Chukchi and Beaufort
Seas and has been reported as far north as 710
24' N.


In Alaska, many commercial fisheries species
and other species are associated with deep
corals. Most associations are believed to be
facultative rather than obligatory. Fish and crabs,
particularly juveniles, use coral habitat as refuge
and as focal sites of high prey abundance. Some
shelter-seeking fishes such as rockfish may use
coral habitat as spawning and breeding sites.

Commercial Fisheries Species Associations
In Alaska, commercial species are managed with
five Fishery Management Plans (FMPs)-Bering
Sea and Aleutian Island (BSAI) Groundfish, Gulf
of Alaska Groundfish, BSAI King and Tanner
Crabs, Salmon, and Scallops. The commercial

Figure 2.13. A shortspine thornyhead (Sebastolobus
alascanus) rests in a field of primnoid gorgonians.
Photo credit: R. Stone, NOAA Fisheries.

harvest of approximately 35 species (or species
groups) is specifically managed with the FMPs.
Most of these species (approximately 85%) are
found during some phase of their life cycle in
deep-water habitats including those inhabited
by deep corals so the potential for associations
between commercial fish species and corals is
high (Figures 2.14 and 2.15).

Heifetz (2002) analyzed data from RACE
survey hauls to determine large-scale (i.e.,
kilometers to tens of kilometers) associations of
commercially targeted fish species with corals.
Rockfish (Sebastes spp.), shortspine thornyhead
(Sebastolobus alascanus), and Atka mackerel
(Pleurogrammus monopterygius) were the
most common fish captured with gorgonians,
scleractinians, and stylasterids. Flatfish
(Pleuronectidae and Bothidae) and gadids were
the most common fish captured with soft corals.

Stone (2006) examined fine-scale (<1 m)
associations of FMP species with corals and
other structure-forming invertebrates from video
footage of the seafloor collected in the central
Aleutian Islands. At the sites surveyed, 84.7%
of the commercially important fish and crabs
were associated with corals and other sedentary
structure-providing invertebrates. All seven
species of rockfish (Sebastes) observed were
highly associated with corals. Associations
ranged from 83.7% for "other" rockfish to 98.5% for
sharpchin rockfish (S. zacentrus). Ninety seven
percent of juvenile rockfish were associated with
corals. Over 20% of the FMP species were in


physical contact with corals and other structure-
forming invertebrates.

Observations from the manned submersible Delta
in the eastern Gulf of Alaska have documented
fine-scale associations (<1 m) of adult shortraker
(S. borealis), rougheye (S. aleutianus), redbanded
(S. babcocki), sharpchin, dusky (S. ciliatus), and
yelloweye rockfish (S. ruberrimus), and golden
king crabs (Lithodes aequispina) with red tree
coral P. pacifica (Krieger and Wing 2002). Large
schools of Pacific ocean perch (Sebastes alutus)
have been observed in dense groves of the
pennatulacean H. willemoesi on the Bering Sea
shelf presumably feeding on dense aggregations
of euphausiids or krill (Brodeur 2001).

Only 16 of the 24 named seamounts in Alaskan
waters summit within the maximum depth range
of FMP species (approximately 3000 m). Several
FMP species have been documented on the
seamounts but studies have not been undertaken
to examine associations of commercial species
and coral habitat. FMP species documented on
Alaskan seamounts include sablefish, longspine
thornyhead (Sebastolobus altivelis), shortspine
thornyhead, rougheye rockfish, shortraker
rockfish, aurora rockfish (Sebastes aurora), and
golden king crabs (Alton 1986; Hughes 1981;
Maloney 2004). Other species of potential
commercial importance found on the seamounts
include the deep-sea sole (Embassichthys
bathybius), spiny dogfish (Squalus acanthias),

Figure 2.14. A darkfin sculpin Malacocottus zonurus rest
a bubblegum coral Paragorgia arborea in one of the seve
gardens surveyed with the submersible Delta. Coral gard
are areas of extraordinary coral abundance and high spec
diversity. Photo credit: R. Stone, NOAA Fisheries.

Figure 2.15. An unidentified eelpout (probably Pu-
zanovia rubra) displays cryptic coloration in a Para-
gorgia colony at 746 m depth in the Aleutian Islands.
Photo credit: R. Stone, NOAA Fisheries.

and several species of grenadiers (Family

Other Species Associations
Many non-commercially important species are
associated with deep corals in Alaska. Both
facultative and obligatory associations are likely
common. Few obligatory associations have
been described to date but recent collections of
micro- and macro-associates of corals should
reveal new examples of unique adaptations and
symbiosis. Forexample, three species ofAleutian
eelpouts (Nalbantichthys sp., Opaeophacus
sp., and Puzanovia sp.) have developed
specializations such as cryptic coloration for life
as adults in Primnoa (Anderson 1994)
and Paragorgia colonies (Figure 2.15).

Observations from the submersible
Delta in the eastern Gulf of Alaska have
documented fine-scale associations
(<1 m) of sea anemones (Cribrinopsis
sp., Stomphia sp., and Urticina sp.),
the basket star (Gorgonocephalus
eucnemis), the crinoid (Florometra sp.),
and the nudibranch (Tritonia exulsans)
with P pacifica (Krieger and Wing 2002).
SAll megafauna were in physical contact
with the coral and were using it as an
.. elevated feeding platform or as refuge.
s., The spiny red sea star (Hippasteria
s under spinosa) was documented preying on
n coral the coral. Macrofauna such as shrimp
ens were also observed within the colonies
cies but were not identified or enumerated.


Macrofaunal assemblages living on deep corals
were studied during the Gulf of Alaska Seamount
cruise in 2004. The chirostylid crab (Gastroptychus
iapsis) and the basket star (Asteronyx sp.) were
the most common macrofauna found on deep
corals (T. C. Shirley, Texas A&M University, pers.
comm.). Other macrofauna collected on corals
included the hippolytid shrimp (Heptacarpus sp.),
actiniarians, crinoids, ophiuroids, crustaceans,
sea stars, pycnogonids, and nudibranchs.
Taxonomic identifications are pending.

Macrofaunal assemblages living on deep corals
collected during the Aleutian Island cruises in
2003 and 2004 were preserved and taxonomic
identifications are underway. Crustaceans,
ophiuroids, and polychaetes appearto be the most
common macro-associates of octocorals (Les
Watling, University of Hawaii, pers. comm.). The
basket star Asteronyx sp. was highly associated
with the deep-sea pennatulacean Anthoptilum
grandiflorum and uses it as an elevated feeding
platform (R. Stone, personal observations).
Many sedentary and sessile taxa are found in
close association with Aleutian Island corals and
include sponges, hydroids, bryozoans, the crinoid
Florometra serratissima, the sea cucumber Psolus
squamatus, and the basket star Gorgonocephalus
eucnemis. More than 100 different species
of sponges, mostly demosponges, have been

Figure 2.16. Calliostomatid snails (genus Otukaia)
prey on the soft flesh of an undescribed species of
bamboo coral at a depth of 1227 m in the central
Aleutian Islands. The snails were recently discov-
ered and are currently being described by Dr. James
McLean at the Natural History Museum of Los Ange-
les County. Photo credit: R. Stone, NOAA Fisheries.

collected during the Aleutian Island studies and
preliminary estimates indicate that more than
200 species of demosponges alone may occur
in association with deep corals in the central
Aleutian Islands (Stone 2006). Sea stars
commonly found in Aleutian Island coral gardens
include Cheiraster dawsoni and Hippasteria
spinosa (R. Stone, unpublished data); the latter
species is a documented predator of octocorals.
Other predators of octocorals include snails of
the genus Otukaia (family Calliostomatidae) that
have recently been observed preying on bamboo
corals in the Aleutian Islands (Figure 2.16).

There are no data regarding commercial fisheries
or non-commercial species associations with
coral habitat in the Arctic region of Alaska.


All known threats to deep coral communities
in Alaska are directly or indirectly the result of
human activities. While activities such as coastal
development, point-source pollution, and mineral
mining have the potential to affect nearshore
habitats, the effects of these activities are
geographically limited and occur or are likely to
occur in areas with minimal coral habitat. Fishing
activities, on the other hand, occur over vast
areas of the seafloor and often in areas containing
sensitive deep coral habitat. Human activities
that may indirectly affect deep coral habitat
include the introduction of invasive species and
changes to the physical and chemical properties
of the oceans due to global warming.

Effects of fishing

Diverse benthic communities on the continental
shelf and upper slope of the Gulf of Alaska,
Bering Sea, and Aleutian Islands support some
of the largest and most important groundfish and
crab fisheries in the world. Alaskan fisheries
within the U.S. EEZ (3 to 200 nm offshore) are
managed under five federal fishery management
plans. Other important fisheries within 3 nm of
shore are managed by the State of Alaska. Four
types of bottom-contact gear are currently used
that potentially affect coral habitat otter trawls,
longlines, pots, and scallop dredges. These
fisheries are distributed from 27 m to about 1000
m, with most effort at depths shallower than


200 m (Stone 2006). The degree to which a
particular gear affects coral habitat depends on
its configuration (i.e., physical area of contact),
operation (i.e., physical forces on the seafloor),
spatial and temporal intensity of operation,
seafloor bathymetry and substratum type,
and the resilience of components of benthic
communities (Table 2.3). Both direct and indirect
effects from fishing activities on corals likely
occur. Direct effects include removal as by-
catch, damage caused by physical contact, and
detachment from the seafloor and translocation
to unsuitable habitat. Indirect effects include
increased vulnerability to predation, especially
for corals detached from the seafloor, and habitat
alteration. Furthermore, there is some evidence
that reproduction is suppressed in damaged
shallow-water scleractinian corals due to a
reallocation of energy reserves for tissue repair
and regeneration (Wahle 1983) and similar effects
may occur in deep non-scleractinian corals.

Disturbance from fishing activities is the greatest
present threat to coral habitat in Alaska (Table
2.3). NMFS estimates that approximately 81.5
metric tons of coral were removed from the
seafloor each year between 1997 and 1999 as
commercial fisheries by-catch in Alaska (NMFS
2004). Approximately 91% of this by-catch
occurs in the Aleutian Islands and Bering Sea
and bottom trawls catch more than 87% of the
total (NMFS 2004). Estimates of the amount of
damaged or detached corals fishing activities
leave behind on the seafloor are not available but
may be substantial. In the centralAleutian Islands,
disturbance to the seafloor from bottom-contact
fishing gear was widespread and approximately
39% of the seafloor on video transects had been
disturbed (Stone 2006). In total, 8.5% of the corals
observed, mostly stylasterids and gorgonians,
were damaged or otherwise disturbed (Stone

Bottom Trawls
Studies worldwide have determined that bottom
trawling alters seafloor habitat and both directly
and indirectly affects benthic communities (Jones
1992; Auster et al. 1996; Auster and Langton
1999; NRC 2002). In addition to removing target
species, bottom trawling incidentally removes,
displaces, or damages non-target species (Ball
et al. 2000), changes the sedimentary properties
of the seafloor (Churchill 1989), and reduces
habitat complexity by physically altering biogenic

structures, including corals, on the seafloor
(Krieger 2001). Such changes can lead to
population level effects on species of economic
importance (Lindholm et al. 1999). Ultimately,
the combination of effects may result in wide-
scale ecosystem change (Gislason 1994; Gori
1998). Directed studies on the effects of bottom
trawling on deep coral habitat in Alaska have
been limited to a few studies (Krieger 2001;
Stone et al. 2005).

Bottom trawls have been extensively used in
Alaskan fisheries since the 1930s. Bottom trawling
has been prohibited east of 1420 W longitude in
the Gulf of Alaska (Figure 2.17A) including the
inside waters of Southeast Alaska, since 1998
but intensive trawling occurred there prior to the
closure. Bottom trawl effort elsewhere in the
state is more continuously distributed (Figures
2.17B-2.17E). Small pockets of intense trawling
for flatfish, Pacific cod, and Pacific ocean perch
have occurred near Kodiak Island in the Gulf of
Alaska (Figure 2.17B) and in the Aleutian Islands
for Atka mackerel and Pacific ocean perch
(Figures 2.17C and 2.17D). NMFS estimates
that approximately 6.2 metric tons of coral are
removed from the seafloor each year by bottom
trawls in the Gulf of Alaska (NMFS 2004). Most
of the Bering Sea has experienced some degree
of exposure to bottom trawls (NMFS 2004) and
several areas have been trawled on average
more than five times per year (Figure 2.17E).

Most bottom trawling occurs on the continental
shelf and upper slope at depths less than 500 m
but some effort does occur to depths greater than

1000 m. Trawling occurs over a wide range of
habitats depending on targeted species and does
occur in areas of coral abundance. Total width
of the trawl system while fishing may reach 110
m, but the area of the seafloor and associated
epifauna contacted by the gear depends on the
design of the otter boards and configuration of
protective gear on other system components
(Stone et al. 2005). Bottom trawling is a major
threat to coral habitat because the area of
seafloor contacted per haul is relatively large, the
forces on the seafloor are substantial, and the
spatial distribution of fishing is extensive (Table
2.3). Areas of the seafloor composed mostly of
bedrock and boulders, and with irregular and
steep bathymetry, are infrequently trawled due to
the risk of damaged and lost gear. Such areas
often support rich coral habitat and may serve as




- 1000m

0 100 200 km

C 0

U'mnalk Island '

S. .. .- , -". ...

.-. '
Boaom Trwl 9-02- 200
o g-. 1000 m

0 00 m200 EEZ
170'W ItW

JD ^ 0

"I Bomers RidgC
u T -'I 9 .

RMtiainTuul J9&^i S. .

,. ... -%. -'" .. -


1000 m

0 100 200 km

Figure 2.17. Average
annual bottom trawl fishing
intensity between 1998
and 2002 in A) the eastern
Gulf of Alaska, B) the west- <.
ern Gulf of Alaska, C) the
eastern Aleutian Islands,
D) the western Aleutian
Islands, and E) the Bering
Sea (adapted from Rose
and Jorgensen 2005).
Trawl intensity is defined in
NMFS (2005), Appendix B.

? Bsonm Trawl 98-02
's ,r'* *

B h.', me.

o limnn
11 1(1 t0 -

de facto sanctuaries from trawl disturbance.

Mid-water Trawls
Mid-water or pelagic trawls are modified bottom
trawls (otter trawls) used to harvest groundfish
near but not on the seafloor. Mid-water trawls
are used exclusively to catch walleye pollock

(Theragra chalcogramma) in the Bering Sea and
are also used in Gulf of Alaska fisheries (see
www.net-sys.com/trawlnets.htm for extensive
descriptions of the various gear used in Alaskan
waters). By regulation, the use of protective
gear on the footrope is not allowed in an effort to
discourage direct contact with the seafloor (NMFS




EIIr 'i~
40 ". "1 '* ', 5



2004). However, the capture of sedentary benthic
species with pelagic trawls is clear evidence that
the gear does make at least occasional contact
with the seafloor. Overall, pelagic trawls likely
have little effect on deep coral habitat in Alaska
since they are seldom fished on-bottom and
typically in areas with minimal coral habitat (Table
Gill Nets
Gill nets are used to harvest Pacific salmon in
estuarine waters of Alaska but are not a threat to
deep coral habitat because they are not used in
areas known to support corals and seldom make
contact with the seafloor.

the hooks (619 of 541,350) fished during the 1998
NMFS longline surveys in the Gulf of Alaska and
Aleutian Islands (Krieger 2001). Longlines may
entangle or hook corals during retrieval (High
1998), while fish attempt to escape during hooking
(R. Stone, personal observations), and dislodge
or damage corals from straining shear during
retrieval (Stone 2006). Derelict longline gear has
been observed entangled in Primnoa colonies
in eastern Gulf of Alaska thickets (R. Stone,
personal observations) and other gorgonians in
Aleutian Island coral gardens (Stone 2006).

A small amount of longline fishing has occurred
on Gulf of Alaska seamounts as evidenced by the

Table 2.3. Potential effects of fishing gears on deep coral habitat in Alaska.

Otter travvls High High High Medium High
Mid-waler Iravls Low Low Medium Low Low
Demersal longlines Medium Low High Medium Medium
Single-set pols Low Medium Medium Low Low
Longline pols High Medium Low Medium Medium
Scallop dredges Medium Low Low Low Low

Bottom Longlines
Longlines are used extensively throughout
Alaskan waters to catch sablefish, Pacific halibut,
Pacific cod, and several species of rockfish to a
depth of at least 1000 m (Figures 2.18A-2.18E).
Bottom (or demersal) longline systems consist of
a mainline to which are attached 1000s of leaders
and hooks (gangions), anchors, and buoyed lines.
Mainlines often stretch 20 km or more across the
seafloor and are often weighted in areas of rough
bathymetry or strong currents. Both ends of the
mainline are weighted with anchors and buoyed
to the surface. No directed studies have been
undertaken to study the effects of longlines on
benthic habitat in Alaska. Longlines are thought
to cause less of an effect on benthic communities
than mobile fishing gear, but by-catch data and
limited in situ observations clearly indicate that
a significant interaction with coral habitat exists.
Longlines are fished extensively in areas of
known coral abundance and by-catch of corals
is common in some areas. For example, corals,
most notably Primnoa, were caught on 0.1% of

recapture of tagged
sablefish there
(Maloney 2004).
Sablefish tagged by
NMFS as part of a
stock assessment
survey have been
recovered by
fishermen on Pratt,
Surveyor, Murray,
Durgin, and Quinn
seamounts in the
Gulf of Alaska.
Scientists believe
that the effort on
the seamounts has

been minimal and has occurred opportunistically
while fishermen transit by the seamounts.

Longlines pose a moderate threat to coral habitat
in Alaska (Table 2.3). They are used extensively
over a broad depth range (Figures 2.18A-2.18E)
and in virtually all habitat types including those
that are typically too rough for trawling. The area
of the seafloor contacted during typical fishing
operations is low but can be more extensive during
gear retrieval in adverse weather conditions.
Straining shear and entanglement are the major
forces on coral habitat and the seafloor. Longlines
are often set in areas of irregular bathymetry
and large arborescent corals such as Primnoa
pacifica, Paragorgia arborea, and black corals
are the most at risk to disturbance.

Pots and Traps
Pots are used extensively throughout much
of Alaska to catch both fish and crabs and are
deployed differently depending on the target
species. Pots are fished singularly for Pacific






200 m k
- 1000 m

0 100 200k

D *'' :..*

^ '
S. Bocr's RidgC

-;:. x

Nulmber .'1 L~me ali. -. ,,- .
..., .,- :":^ ^
,, -'+ .+ .,%

O 100 200 km

Figure 2.18. Number of
longline hauls between
1973 and 1996 in A) the
eastern Gulf of Alaska, B)
the western Gulf of Alaska,
C) the eastern Aleutian
Islands, D) the western
Aleutian Islands, and E) the
Bering Sea (Adapted from
Fritz et al. 1998). Method-
ology described in Fritz et
al. 1998.

cod and sablefish in the Gulf of Alaska (Figures
2.19A and 2.19B) and additionally for Greenland
turbot (Reinhardtius hippoglossoides) in the
Aleutian Islands (Figures 2.19C and 2.19D)
and Bering Sea (Figure 2.19E). Pot fishing is
typically highly localized in these areas (Figures
2.19A-2.19E). Important fisheries with single

pots for king crabs (Paralithodes camtschaticus,
P. platypus, Lithodes aequispina), Tanner crabs
(Chionoecetes bairdi), snow crabs (Chionoecetes
opilio), and Dungeness crabs (Cancer magister)
occur in the Gulf of Alaska and Bering Sea. Pot
fisheries also occur for golden king crabs in the
Aleutian Islands (Figures 2.19A and 2.19B). In

Balhymetry k
2- 00 m




0 100 200kr

C 0

-f- e
I~mnuklail~Ji C-.

2- 00M
Number 6f Pot Hauls


0 100 200 kEEZ
17CW 160W


Bower's Ridge

iinter ni Poi Hauk -


B a l 'y" e r y

0 100 200

Figure 2.19. Number of
fish pot hauls between
1973 and 1996 in A) the
eastern Gulf of Alaska, B)
the western Gulf of Alaska,
C) the eastern Aleutian
Islands, D) the western
Aleutian Islands, and E)
the Bering Sea (Adapted
from Fritz et al. 1998).
Methodology described in
Fritz et al. 1998.


>.-' -



this fishery, however, pots are strung together in
strings of 10 to 90 pots or more and total weight
of the gear per string can exceed 30 metric tons.
Pots are strung together with 1-inch or larger
diameter polypropylene line and a single longline
may stretch between 3 and 9 km. The fishery
occurs at depths between 100 and 719 m and in

a wide range of habitats on the slope, offshore
banks, and offshore pinnacles that include rocky
areas with irregular bathymetry.

No studies have been undertaken to study the
effects of pot fishing on seafloor habitat in Alaska.
Single pot fisheries likely have a minimal effect on



... ~

- -

- :jj,:I


Umnak Island

NuiLwr of Crab Nis

SBowel's Ridge

0 100 200la 1000m

the golden king crab (Lithodes aequispina) fishery be-
nds and B) the western Aleutian Islands. Each area re
)urce data: Alaska Department of Fish and Game.

t- the strength limitations of the longline; howe
d under certain conditions the gear can be drag
is like a plough across the seafloor. This situa
s can occur in areas of steep bathymetry and w
b strong winds and currents dictate that fisl
e vessels retrieve gear while being forced a
3) from it. At one site in the central Aleutian Isl
,e where disturbance from this gear was obser
s with the submersible Delta (Figure 2.21),
ar seafloor was scoured to bare substrate alone(
ly strips (Stone 2006). Aleutian Island coral gard
>r are at high risk to disturbance from this fisher
:o Scallop Dredges
A small but important fishery has occurred
the weathervane scallop Patinopecten caure
in the Gulf of Alaska and Bering Sea since 1
(Shirley and Kruse 1995). The fishery occur
relatively well-defined areas of unconsolidd
soft sediments on the continental shelf
at depths between 60 and 140 m (Turk 2(
Barnhart 2003). Scallop dredges are drag
along the seafloor and designed to dig into
top layer of sediment. Dredges have a maxin
width of 4.6 m. No directed research on
effects of scallop dredges on coral habitat
been undertaken in Alaska. Overlap does ob
between the fishery and the known distribution
pennatulaceans, including ecologically impor
pennatulaceans, including ecologically impor


*ibution of fishing is small and the area of the pi
loor contacted per tow is relatively small. is
le the gear is intrusive, it is generally used re
oft-sediment areas where coral abundance
)w. Groves of pennatulaceans in the Gulf N
laska are most at risk to disturbance from di
lop dredges. w
'cts of Other Human Activities A
and Gas Exploration and Extraction fr
hore oil and gas operations in Alaska include rr
oration, development, and production tl
/ities (NMFS 2005). Most of these activities Ic
;ently occur in Cook Inlet in the Gulf of Alaska in
on the North Slope (Beaufort and Chukchi c.
s)-areas of Alaska that do not support c.
ificant deep coral communities. Disturbances al
1 these activities that may affect coral habitat h;
ide physical alterations to habitat, waste tl
charges (well drilling cuttings and muds), o
oil spills. Cimberg et al. (1981) discuss rr
potential effects of oil and gas development in
Jeep corals in Alaska. They concluded that u:
skan corals are unlikely to suffer adverse tl
cts from oil and gas development, because cc
t of the known deep coral distributions do not tl
jr in lease areas and areas where platforms re

t physiologically sensiti\
) corals, the planula lan
ersal, and therefore ur

e life history stage of IV
'al stage, is brief and cc
likely to be affected e:

arch has been undertaken to study the effects D
il toxicity on any life history stage of deep a
Is found in Alaska. The potential for effects A
these activities on coral habitat is likely to tl
.ase in the future as the world's demand for
nd gas products continues to increase. P
foyment Of Petroleum Pipelines And rr
imunication Cables di
k Inlet is the only area of the state where al
aleum pipelines (specifically crude oil) have o0
i deployed in benthic marine habitats. bi
teen pipelines totaling 141 km in length were di
oyed on the seafloor of Cook Inlet between g
5 and 1986 (Robertson 2000). Eight state and di
ral agencies have regulatory authority over ol
lines in Cook Inlet. Accidental spills have di
irred in the past and are likely to occur in the ol
*e as many of the pipelines reach the end of pl
expected life span (Robertson 2000). The ol

lines has not been inventoried but the region
,t known to support abundant or diverse coral

ierous communication cables have been
oyed on the ocean floor throughout Alaskan
irs since 1900. Thousands of kilometers
ables stretch along the seafloor between
kan communities and ports in Washington
Oregon. Cables have been deployed
I the shoreline down to depths of 7000
There are no known regulations governing
placement of submarine cables but their
tions are accurately mapped so that potential
'actions with other seafloor uses (e.g., fishing)
be avoided. There are no known reports of
e deployments directly affecting coral habitat
)ugh there is some likelihood that cables
) been placed in coral habitat, especially in
Aleutian Islands. Cables are typically laid
he seafloor where they remain exposed but
be buried using specially designed ploughs
areas where bottom fishing and other seafloor
occur. In areas where cables are exposed
may provide attachment substrates for
Is and other emergent epifauna and may
before provide a known time-line for studies of
jitment and subsequent growth of emergent
auna that settle on them (Levings and
)aniel 1974). No such studies have been
jucted yet in Alaska but clearly the potential
ts to use submarine cable deployments to
insights into coral habitat recovery rates.
loyment of communication cables is presently
inimal threat to deep coral ecosystems in
ka given the very small area of the seafloor
is contacted by them.

jtion Point-source Discharges
it-source discharges that occur in coastal
ne areas ofAlaska have little potential to affect
Scoral habitat. Coral habitatissparseincoastal
is of Alaska where point-source discharges
ir or are expected to occur in the near future
a few coastal areas near municipalities
support groves of pennatulaceans. The
test threat to coral habitat from point-source


also cause algal blooms that are lethal to corals
(Alcolado 1998). Chlorine is toxic to marine life,
and chlorinated sewage effluent may subject
marine biota, including octocorals, to eithersingle-
event acute exposures or to chronic exposures
(Tomascik et al. 1997).

Fish Processing Waste
I nAlaska, seafood-processing facilities are located
both on shore and at sea onboard processing
vessels. Coral habitat is sparse in coastal areas
of Alaska where seafood-processing discharges
occur and concerns to coral habitat there would
be similar to those for point-source discharges.
At-sea processors would have little effect on deep
coral habitat unless they routinely discharged
waste in areas of high coral abundance.

Harvest of Precious Corals
A directed fishery for precious corals never
developed in Alaska despite the fact that several
species have potential commercial value as
jewelry (Cimberg et al. 1981). Corals found in
Alaska with potential commercial value include
Primnoa pacifica, Primnoa wingi, bamboo corals
(Family Isididae), black corals, and a single
species of precious red coral (Family Coralliidae)
reported from Patton Seamount in the Gulf of
Alaska (A. Baco-Taylor, WHOI, pers. comm.).
However, many corals that are collected as by-
catch, particularly P pacifica, bamboo corals,
and stylasterids, are often retained by fishermen
as souvenirs and curios.

Mineral Mining
Mineral mining operations in Alaska have been
limited to offshore placer mining for gold and
barite off the coast of Nome in Norton Sound
(northern Bering Sea) and at a single location
near Petersburg in Southeast Alaska (Conwell
1976). Mineral mining activities could potentially
affect deep coral habitat through increased
sedimentation and turbidity near the seafloor but
are unlikely to occur in areas of coral abundance
in the near future.

Climate Change
Climatic regime shifts and cyclic environmental
fluctuations associated with Pacific Decadal
Oscillations, El Nilo/Southern Oscillation Climate
and La Nina events have had documented effects
on oceanographic and biological processes
in the North Pacific Ocean. Effects on corals
of this interannual to decadal variability have

not been reported. Long-term climatic change
due to global warming could affect seawater
temperature, salinity, density, sea level, and
ambient light levels especially in shallow and
nearshore waters. None of these changes is
expected to cause direct mortality of deep corals
or significantly alter their geographic or depth
distribution but effects on growth rates and food
supply (i.e., phytoplankton) are possible.

Increases in atmospheric carbon dioxide caused
by manmade emissions have been linked to
decreases in oceanic pH (Caldeira and Wickett
2003). Decreases in oceanic pH and resulting
decreases in calcium carbonate saturation
state and calcification could have devastating
effects on marine organisms dependent on the
extraction of calcium carbonate from seawater for
skeletal building (Kleypas et al. 1999; Guinotte
et al. 2006). Zooxanthellate corals in shallow
waters will experience decreasing aragonite
saturation states that could negatively affect
their calcification rates and the stability of reef
ecosystems (Guinotte et al. 2003). Numerous
studies have shown substantial decreases in
calcification rates (>40%) with relatively modest
decreases in aragonite saturation state (Langdon
et al. 2003; Langdon and Atkinson 2005). Some
evidence suggests that deep-sea biota may be
sensitive to changes in pH (Seibel and Walsh
2001; Guinotte et al., 2006; Roberts et al., 2006).
Mounting evidence suggests that if CO2emissions
continue as projected, undersaturated regions
will develop in the sub-arctic and polar regions of
Alaska by the end of the 21st century (Orr et al.
2005; Kleypas et al. 2006; Guinotte et al. 2006).
Scleractinian corals would be most affected by
this development, but are not important structure-
forming corals in Alaskan waters. Octocorals,
stylasterids and pennatulaceans however, are
important structure-forming components of
benthic ecosystems in Alaskan waters and will
likely be affected by ocean acidification. The
sclerites of octocorals are calcitic, but the axes
may be composed of calcite, aragonite, or
amorphous carbonate hydroxylapatite (Bayer and
Macintyre 2001). The calcite saturation horizon,
along with the aragonite saturation horizon, is
moving to shallower depths over time (Feely et
al. 2004), which could affect all corals in Alaska
that use calcite to build skeletal tissue.

Invasive Species
The introduction of invasive species to Alaskan


waters is a real threat and the State of Alaska
has developed an Aquatic Nuisance Species
Management Plan to prevent introductions and
identify and respond to threats (ADF&G 2002).
Ballast water discharges from ships and barges
are the single largest potential source of invasive
species in Alaska. For example, tankers arriving
from domestic ports at Port Valdez, Prince
William Sound, release the third largest volume
of ballast water of all U.S. ports (ADF&G 2002).
Tankers arriving from foreign ports are required to
exchange ballast water at sea (in waters at least
2000 m deep). The potential for introductions
in coastal Alaska and the Aleutian Islands in
particular is high given the high volume of ship
traffic from ports around the world.

To date, the introduction of invasive species has
been largely limited to a few species of freshwater
fish and aquatic plants. There are no known
invasive species of corals or predators of corals in
Alaskan waters although the threat of introduction
exists. The threat of introduction may increase
if more favorable oceanic conditions related to
climatic change develop in the future.


The North Pacific Fishery Management Council
(NPFMC) manages the fishery resources of
Alaska with five Fishery Management Plans
(FMPs). The Magnuson-Stevens Fishery
Conservation and Management Act (MSFCMA)
mandates that FMPs must include a provision to
describe and identify essential fish habitat (EFH)
for each fishery, minimize to the extent practicable
adverse effects on such habitat caused by fishing,
and identify other actions to encourage the
conservation and enhancement of such habitat.
EFH has been broadly defined by the Act to
include "those waters and substrate necessary to
fish for spawning, breeding, feeding, or growth to
maturity." Deep coral habitat has been identified
as EFH for some groundfish species (Witherell
and Coon 2001) and several areas of Alaska
have recently been designated as Habitat Areas
of Particular Concern (HAPCs) and are presently
afforded some protection from disturbance
by fishing activities (described below). The
Minerals Management Service (US Department
of the Interior) oversees petroleum and mineral
resource development in the offshore waters of

the U.S. EEZ and implements studies designed
to predict the effects of resource development
on the marine ecosystem including deep coral

Seafloor Mapping
Approximately 46,710 km2 of seafloor habitat
has been mapped in the Alaska region using
multibeam sonar technology (Table 2.4). These
efforts have been piecemealed together by several
agencies including NMFS, Alaska Deparment
of Fish and Game (ADF&G), National Park
Service (NPS), University of Alaska Fairbanks,
and Oregon State University. Additionally, about
27,780 km2 of seafloor has been mapped by
NOAA's National Ocean Service (NOS) since
1994 for navigational purposes. No coordinated
plan to map the seafloor within the EEZ currently
exists and mapping efforts to date have been
scattered from Southeast Alaska through the
Aleutian Islands including some of the seamounts
within the EEZ. Mapping has included 4,220 km2
and 28,280 km2 of seafloor on Gulf of Alaska
shelf and slope habitats and Gulf of Alaska
seamounts, respectively. An additional 14,150
km2 of seafloor has recently been mapped in the
Aleutian Islands.

While the purpose of the seamount and some of
the Aleutian Island mapping efforts have been
strictly to support detailed studies on deep coral
habitat, most of the mapping efforts to date have
been in support of studies on essential fisheries
habitat and geological processes. Additional
goals of these studies have been to determine
the effects of fishing on benthic habitat, fish stock
assessments, understanding basic ecological
processes, and life history studies of benthic
organisms (e.g., Shotwell etal. in press; O'Connell
et al. in press).

Several of the mapping efforts have included
the collection and subsequent interpretation of
backscatter data and the detailed classification of
seafloor habitats using the methods of Greene et
al. (1999). Direct observations of the seafloorwith
occupied submersibles, ROVs, or towed cameras
have been used to ground-truth habitat types and
provide fine-scale resolution of habitat features.
One goal of the Aleutian Island studies (see http://
listing.htm) is to develop a model to predict
where deep coral habitat is located throughout
the region. Mapped areas were systematically


Table 2.4. Areas of Alaska that have been mapped with modern multibeam sonar technology. Al =
Aleutian Islands, GOA = Gulf of Alaska, SM = seamounts, UAF = University of Alaska Fairbanks,
NMFS = National Marine Fisheries Service, ADFG = Alaska Department of Fish and Game, OSU =
Oregon State University, NPS = National Park Service.

GOA Hazy Islands ADFG 50-300 Yes 390
GOA Cape Ommaney ADFG, NMFS 30-300 Yes 275

GOA Glacier Bay National Park NPS 10-410 Yes 62

GOA Fairweather Ground ADFG 23-192 Yes 280
GOA Yakutat Bay ADFG 15-50 Unknown 20
GOA Yakutat Bay ADFG 15-50 Unknown 20
GOA South Yakutat NMFS 190-1045 Unknown 372
GOA Pamplona Spur NMFS 120-940 Yes 162
GOA Portlock Bank NMFS 100-750 Yes 790
Albatross Bank,
GOA Snakehead NMFS 60-810 Yes 310
Albalross Bank, 8-fathom
GOA pinnacle NMFS, NOS 20-716 No 17
Albatross Bank, 49-
GOA fathom pinnacle NMFS 80-800 Unknown 32
GOA Chirikof UAF 100-600 Unknown 1,550
GOA. SM Seamounts 2002 UAF, OSU ? Yes ?
GOA. SM Seamounts 2004 UAF, OSU ? Yes 14,081
Transit between
GOA, SM seamounts UAF, OSU ? Unknown 9,000

GOA, SM Derickson Seamount UAF, OSU 2750-6800 Yes 5,200
Al Akutan UAF 78-482 Unknown 27
Al Bogoslof UAF 20-820 Unknown 28
Al RV Revelle transit UAF 90-4200 Yes 11,341

Al Samalga Island, North NMFS 107-323 Yes 11

Al Samalga Island, South NMFS 120-150 Yes 9
Islands of Four
Al Mountains, North NMFS 144-223 Yes 13
Islands of Four
Al Mountains, South NMFS 88-204 Yes 12
Islands of Four
Al Mountains, West NMFS 116-218 Yes 11
Al Aleutian Corals NMFS 100-3000 Yes 2,697
Al Track Lines NMFS 30-4000 Yes NA
Total 46,710


selected so that results can be extrapolated to
unmapped areas. Habitats within the mapped
areas are currently being classified through
interpretation of the bathymetric and backscatter
data. Submersible and ROV observations are
being used to ground-truth the habitat types,
map coral observations, and ultimately to provide
data on coral densities relative to mapped habitat

Ongoing Research
Research activities in 2006 focused on completing
taxonomic, genetic, and reproductive ecology
analyses on more than 400 coral specimens
collected during the 2003-04 Aleutian Island
studies and 140 coral specimens collected
during the 2004 Gulf of Alaska seamount cruise.
Additionally, detailed examination of video
footage collected from submersibles and ROVs
during these studiesis underway and willto
provide fine-scale data on coral distribution,
habitat characteristics, species associations,
and disturbance from both human and natural

The submersible Delta was used in 2005 to
delineate the extent of Primnoa thickets in two
areas of the eastern Gulf of Alaska (Fairweather
Ground and Cape Ommaney; Figure 2.22). The
two areas were established as HAPCs by NMFS
in July 2006 and the use of all bottom-contact
fishing gear is now prohibited in those areas.
The purpose of the research was to provide
detailed data on the distribution of Primnoa in
the areas so that the efficacy of the closures to
protect the thickets from incidental disturbance
can be predicted. Additional objectives of the
research are to assess the present condition
of the thickets, examine the fine-scale use of
the coral habitat by FMP species, and collect
specimens for taxonomic identification. A third
site in Dixon Entrance near the maritime boundary
with Canada was also investigated to determine
if the Primnoa thickets reported from that region
(Krieger 2001) warrant designation as a HAPC.
The thickets in that region appear to be located in
deeper water and in a region where both the U.S.
and Canada claim jurisdiction. A joint research
cruise by both governments may be planned in
the future to examine coral habitat in that region.

A two-year study to examine shallow-water
populations of Primnoa pacifica in Glacier Bay
National Park was completed in April 2005.

Populations of Primnoa were discovered in
2004 along bedrock walls recently exposed by
retreating glaciers (Figure 2.23). The study is
investigating the role of oceanographic processes
in coral depth distribution and the potential use
of an accurate deglaciation record to validate
estimated growth rates for the species (Stone
et al. in preparation). Thriving populations of
Primnoa were discovered in two additional glacial
fjords in Holkham Bay, SoutheastAlaska during a
research cruise in 2006. Samples were collected
from 80 colonies from four spatially distinct
"populations" during a second cruise to the fjords
in 2007. Those samples will be used to develop
microsatellite genetic markers to examine the
population structure of Primnoa in the fjords and
to address questions regarding larval dispersal
and gene flow.

Planned or Anticipated Activities
No directed research or mapping activities are
planned at the present time due to limited funding
and personnel support. Several important areas
of deep coral research remain a high priority
for the region (discussed below) and those will
be addressed as funding and support becomes
available in the future.

North Pacific Fishery Management Council
The North Pacific Fishery Management Council
has responsibility for developing fishery
management plans for the nation's groundfish
resources in the EEZ of the Alaska region and
oversees the implementation of measures to
conserve and enhance essential fish habitat for
those resources. Since 1987, 1,107,890 km2 of
seafloor habitat in the Bering Sea and Aleutian
Islands has been afforded some protection from
fishing activities (Figure 2.22). An additional
202,380 km2 of seafloor habitat has been
afforded some level of protection in the Gulf of
Alaska (summarized in NMFS 2004). Most area
closures are for specific gear types only, others are
seasonal, and some closures go into effect only
when a species by-catch cap has been reached.
Year-round closures to trawl gear exist in both the
Bering Sea and Gulf of Alaska to protect important
crab habitat. Year-round closures also exist
around Steller sea lion rookeries to protect forage
species. A single area, the 8.1 km2 Edgecumbe
Pinnacles Marine Reserve (also known as the
Sitka Pinnacle Marine Reserve) in the eastern
Gulf of Alaska, was established in 2000 as the
only no-take groundfish reserve in the state. A

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