The State of 

Deep Coral Ecosystems of 

the United States: 2007

Produced by the National Oceanic and 

Atmospheric Adminstration

NOAA TEcHnical Memorandum CRCP - 3
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 Memorandum 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/

Disclaimer:

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.

i
The State of Deep Coral Ecosystems

of the United States

Report Editors:

S. Elizabeth Lumsden

Thomas F. Hourigan

Andrew W. Bruckner

Gabrielle Dorr

National Oceanic and Atmospheric Administration

October 2007

NOAA Technical Memorandum CRCP 3

United States Department 

of Commerce

Carlos M. Gutierrez

Secretary

National Oceanic and 

Atmospheric Administration

Conrad C. Lautenbacher, Jr.

Administrator

National Marine Fisheries 

Service

William T. Hogarth

Assistant Administrator

ii
Acknowledgements

Overall Report and Chapter 1: Introduction and Overview

The editors would like to thank Michael Peccini for developing GIS images and numerous others who provided 

photographs and images for use in this chapter. We would also like to thank the many reviewers for contributing considerable time and effort to provide constructive comments on this chapter. An independent external 

peer review of the full report was conducted by the Center for Independent Experts (CIE) at the University of 

Miami. Reviewers for the CIE review included Stephen Cairns – Smithsonian Institution, J. Anthony Koslow 

– Scripps Institution of Oceanography, and Pål B. Mortensen – Institute of Marine Research, Bergen, Norway. 

The report also benefi ted from additional external reviews by the South Atlantic Fishery Management Council, Western Pacifi c Fishery Management Council, New England Fishery Management Council, Peter Auster 

– NURC University of Connecticut, Andrew Shepard – NURC University of North Carolina, Wilmington, Chris 

Kelley – Hawaii Undersea Research Laboratory, Robert Y. George --George Institute for Biodiversity and Sustainability, John K. Reed – Harbor Branch Oceanographic Institution, Peter Etnoyer – Harte Research Institute, 

Texas A&M, John Warrenchuck and Santi Roberts – Oceana, Lance Morgan and Fan Tsao – Marine Conservation Biology Institute, Alberto Lindner – Smithsonian Institution, Brian Tissot – Washington State University. 

Internal NOAA reviewers included: NOS comments from Ed Bowlby – OCNMS, Mary Sue Brancato --OCNMS, 

Emma Hickerson – FGBNMS, G.P. Schmahl – FGBNMS, Roger Griffi s and Kara Meckley --CRCP, Steve Gittings and Brad Barr – NMSP, Jeff Hyland – NCCOS, OAR Comments from NURP and OE, NMFS comments 

from Southeast Fisheries Science Center, Southeast Regional Offi ce, Northeast Fisheries Science Center, 

Northwest Regional Offi ce, Southwest Fisheries Science Center, Alaska Regional Offi ce, Dwayne Meadows 

– Offi ce of Protected Resources, Robert Brock – Offi ce of Science and Technology. The Introductory Chapter was a collaborative effort with the authors from each of the regional chapters, and their input has greatly 

strengthened this section.

Chapter 2: Alaska Chapter 

Jon Heifetz (AFSC) provided the overview on multi-beam mapping efforts in Alaskan waters and reviewed 

an earlier draft of this chapter. Jana DaSilva Lage and Rob Hansen (Fugro Pelagos, Inc.) provided helpful 

information regarding submarine telecommunication cable deployments in Alaskan waters. Amy Baco-Taylor 

(Woods Hole Oceanographic Institution), Peter Etnoyer (Texas A&M University), and Tom Shirley (Texas A&M 

University) provided information on coral distribution from the Gulf of Alaska seamounts. John Guinotte (Marine Conservation Biology Institute) provided helpful insights regarding the effects of ocean acidifi cation on 

North Pacifi c corals. Jennifer Reynolds (University of Alaska Fairbanks) provided information about the submarine geology of the North Pacifi c Ocean. Cathy Coon (NPFMC) provided detailed information on fi shing 

area closures in Alaskan waters and Jon Warrenchuk (Oceana) provided Figure 2.13. 

Chapter 3: West Coast

Numerous agencies, institutions and individuals provided data and input to Chapter 3, including Mary Yoklavich 

(NMFS, Southwest Fisheries Science Center), Brian Tissot and Jennifer Bright (Washington State Univ. Vancouver), Milton Love (Univ. California, Santa Barbara), Ed Bowlby and Mary Sue Brancato (NMSP, Olympic 

Coast National Marine Sanctuary), Jeff Hyland (NOS, National Centers for Coastal Ocean Science), Jodi Pirtle 

(formerly at Washington State Univ. Vancouver), Dan Howard and Dale Roberts (NMSP, Cordell Bank National 

Marine Sanctuary), Erica Burton (NMSP, Monterey Bay National Marine Sanctuary), Mark Wilkins (NMFS, 

Alaska Fisheries Science Center), Lance Morgan (Marine Conservation Biology Institute), Peter Etnoyer 

(Aquanautix Consulting), Chris Goldfi nger, Chris Romsos and Mark Hixon (Oregon State Univ.), Gary Greene 

(Moss Landing Marine Laboratories), Rikk Kvitek (California State Univ. Monterey Bay), Alberto Lindner (formerly with Duke Univ.) and Glen Jamieson (Department of Fisheries and Oceans Canada).

In addition to reviews by the Center for Independent Experts, additional reviews of all or portions of Chapter 3 

were provided by Waldo Wakefi eld and Ewann Berntson (NMFS, Northwest Fisheries Science Center), Mary 

Yoklavich (NMFS, Southwest Fisheries Science Center), Ed Bowlby and Mary Sue Brancato (NMSP, Olympic 

Coast National Marine Sanctuary), Jeff Hyland (NOS, National Centers for Coastal Ocean Science), Frank 

iii
Lockhart (NMFS Northwest Regional Offi ce), Steve Copps (NMFS, Northwest Regional Offi ce) and Lance Morgan and Fan Tsao (Marine Conservation Biology Institute). We appreciate all their constructive comments. 

Chapter 4: Hawaii 

Much of the research reported in chapter 4: the Western Pacifi c Region: Hawaii and the US Pacifi c Islands, 

was supported by the NOAA Offi ce of Ocean Exploration and the NOAA Undersea Research Program through 

the Hawaii Undersea Research Laboratory. Stephen Cairns and Dennis Opresko provided preliminary identifi cations and unpublished species lists to help us provide as complete a taxonomic inventory as possible. 

We are also grateful to Celeste Mosher, Deborah Yamaguchi and Ronald Hoeke who helped with tables and 

graphics. 

Chapter 5: Northeast

The authors thank Beth Lumsden, Tom Hourigan, and other members of the Deep Coral Team for leading 

this effort and for editing and formatting our chapter. The authors would also like to thank reviewers Stephen 

Cairns, Pål Mortensen, and J. Anthony Koslow, as well as additional reviewers, for constructive comments and 

suggestions.

Chapter 6: Southeast

We thank the NOAA Offi ce of Ocean Exploration, the U.S. Geological Survey, the South Atlantic Fishery 

Management Council (SAFMC), Environmental Defense, and the Minerals Management Service for helping to 

fund our deep coral research, which contributed to this review. Much of these data were collected as part of 

a team effort involving the authors, K.J. Sulak (USGS), E. Baird (NC Museum of Natural Sciences), C. Morrison (USGS) and A. Howard. Andy Shepard (National Undersea Research Center, UNC-Wilmington) facilitated several of our projects, including this report, and provided the Oculina photographs. We acknowledge 

the efforts of the SAFMC in leading the way toward better management of deep coral habitats. We thank A.M. 

Quattrin and M.L. Partyka for help with fi gures and data analysis. This chapter was partially supported by the 

NOAA Ecosystem Assessment Division.

Chapter 7: Gulf of Mexico

The authors would like to extend their appreciation and thanks for all the contributions from reviewers, which 

have greatly improved this chapter. A special ‘Thank You’ goes to Dr Steven Cairns (National Museum of Natural History) who helped us navigate the complexities of coral taxonomy, and to the staff of the Flower Garden 

Banks National Marine Sanctuary who generously shared their research with us’

Chapter 8: Caribbean

The authors would like to thank the following: Stephen Cairns, of the National Museum of Natural History, 

who’s foundation work, advice, patience, and comments were invaluable; Nancy Voss, who helped us through 

the collections at the University of Miami’s Invertebrate Museum; Loretta Burke, of the World Resources Institute, for our baseline bathymetry map; John Reed, Judith Lang, and Charles Messing, who provided fi rst-hand 

accounts of lithoherms in the Straits of Florida; Daniel Opresko, who’s expertise with deep water black corals 

brought us into the light; and many others. Support was provided by Marine Conservation Biology Institute and 

the University of Miami’s Coral Reef Conservation Research Laboratory. Special appreciation goes to the late 

Peter Lutz, for advice, counsel, and support.

iv
The State of the Deep Coral

Ecosystems of the United States

NOAA Technical Memorandum CRCP-3

Table of Contents

Acknowledgements iii

Table of Contents v

Preface vi

Chapter 1: Deep Coral Ecosystems of the United States: Introduction and National Overview

Thomas F. Hourigan, S. Elizabeth Lumsden, Gabrielle Dorr, Andrew W. Bruckner, 

Sandra Brooke, Robert P. Stone 1

Chapter 2: State of the U.S. Deep Coral Ecosystems in the Alaska Region: Gulf of Alaska, Bering 

Sea and the Aleutian Islands

 Robert P. Stone and S. Kalei Shotwell 65

Chapter 3: State of the U.S. Deep Coral Ecosystems in the United States Pacific Coast: California 

to Washington

Curt E. Whitmire and M. Elizabeth Clarke 109

Chapter 4: State of the U.S. Deep Coral Ecosystems in the Western Pacific Region: Hawaii and the 

United States Pacific Islands 

 Frank A. Parrish and Amy R. Baco 155

Chapter 5: State of the U.S. Deep Coral Ecosystems in the Northeastern United States Region: 

Maine to Cape Hatteras 

 David B. Packer, Deirdre Boelke, Vince Guida, and Leslie-Ann McGee 195

Chapter 6: State of the U.S. Deep Coral Ecosystems in the Southeastern United States Region: 

Cape Hatteras to the Florida Straits

 Steve W. Ross and Martha S. Nizinski 233

Chapter 7: State of the U.S. Deep Coral Ecosystems in the Northern Gulf of Mexico Region: Florida 

Straits to Texas 

 Sandra Brooke and William W. Schroeder 271

Chapter 8: State of the U.S. Deep Coral Ecosystems in the United States Caribbean Region: 

Puerto Rico and U.S. Virgin Islands 

 Steven J. Lutz and Robert N. Ginsburg 307

v
PREFACE

This report represents the first effort by the National Oceanic and Atmospheric Administration (NOAA), 

in partnership with other federal, academic and non-governmental partners, to bring together available 

information on the abundance and distribution of structure-forming corals that occur in U.S. waters at 

depths greater than 50 m. It consists of an introduction, National Overview and seven regional chapters 

describing deep coral communities in U.S. waters off Alaska, the U.S. West Coast, Hawai’i and the U.S. 

Insular Pacific, the Northeastern U.S., Southeastern U.S., Gulf of Mexico, and U.S. Caribbean. This 

report reflects the tremendous increase in awareness of these communities that has evolved over the 

last few years as the result of increasing exploration and research to understand deeper regions of the 

oceans. In the U.S., NOAA is proud to serve as a leading partner in much of this work.

NOAA coordinated the development of this report, under the auspices of the Deep Coral Team of the 

NOAA Coral Reef Conservation Program. It reflects the work and dedication of writing teams from each 

region and these teams should be cited as primary authors of the regional chapters. The report also 

benefited from the comments and suggestions of numerous federal and external reviewers and a Data 

Quality Act peer review coordinated through the Center for Independent Experts. 

An introductory chapter defines and provides background information on structure-forming deep corals 

and identifies major threats that they face. A National Overview explores general trends in these 

communities across the regions from a national perspective. Chapters 2 through 8, the regional chapters, 

were developed by authors considered experts in the field of deep coral research and management 

and those chapters represent the core of this report. The authors of each chapter briefly describe the 

region and geological and oceanographic features important to deep coral communities; identify the 

major deep coral taxa that structure habitats in the region and what is known about their distribution; 

provide information on the other species associated with coral habitat; describe the threats to these 

habitats; discuss management efforts developed to respond to these threats, and briefly outline regional 

information needs.. The report also includes unpublished data and observations collected during recent 

research expeditions.

This report fulfills a commitment made in the U.S. Ocean Action Plan as part of an overall effort to 

research, survey and protect deep coral communities. It reflects NOAA’s growing understanding of 

the importance of these communities as hot-spots for deep-water biological diversity, and NOAA’s 

commitment to ensuring their enhanced conservation. This report is also a central part of a broader 

NOAA effort to develop a National Deep Coral and Sponge Research, Conservation and Management 

Strategy. We hope that this first Report on the State of Deep Coral Ecosystems of the United States

will stimulate additional research, surveys and protection, and hope that periodic future reports will 

document both increased understanding and protection of these unique and valuable ecosystems. 

vi

STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES: 

INTRODUCTION AND NATIONAL OVERVIEW

Thomas F. Hourigan1, S. Elizabeth Lumsden1, Gabrielle Dorr2, Andrew W. Bruckner1, 

Sandra Brooke3, Robert P. Stone4

1

NOAA National Marine Fisheries Service, 

Office of Habitat Conservation 

1315 East West Hwy SIlver Spring, MD 20910

2

NOAA National Marine Fisheries Service, 

Southwest Regional Office

3

Ocean Research and Conservation Association, 

Fort Pierce, Florida 34949

4

Auke Bay Laboratory, National Marine 

Fisheries Service, Alaska Fisheries Science Center, 

11305 Glacier Highway, Juneau, Alaska 99801-8626

Figure 1.1 An Alaskan “coral garden” with several 

species of soft corals, hydrocorals, hydroids, and 

demosponges. Photo credit: Alberto Lindner

INTRODUCTION

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 

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 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

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 occurring 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 neutral 

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 

fishes.

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, slowgrowing 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. 

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 

sparse. 

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 

WHAT ARE STRUCTURE-FORMING 

DEEP CORALS?

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. Structureforming 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, 

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 Linné wrote 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

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 structureforming deep corals. 

PHYLUM CNIDARIA

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)

III. Class Cubozoa - does not contain corals

IV. Class Scyphozoa - does not contain corals

1

Taxonomic 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.)

4

Not all taxonomists recognize the order Gorgonacea as separate from Alcyonacea.

5

The 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.).

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 structureforming deep corals. 

PHYLUM CNIDARIA

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)

III. Class Cubozoa - does not contain corals

IV. Class Scyphozoa - does not contain corals

1

Taxonomic 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.)

4

Not all taxonomists recognize the order Gorgonacea as separate from Alcyonacea.

5

The 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.).

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 nonreef-building corals. Although only a few stony 

coral species (order Scleractinia) form deepwater 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. 

2006). 

Shallow corals need well-known and welldocumented 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 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

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

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

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

Parameter Tropical shallow stony corals1 Deep stony corals1

Depth range 0-100 m 39-3,000 m

Temperature 18-31o C 4-13o C

Distribution Tropical and subtropical seas from 

30oN-30oS

Potentially global, at least 

56o S-71o N

Symbiotic Algae Yes No (Note: several species of Oculina 

and Madracis have a facultative 

relationship with zooxanthellae in 

shallow populations)

Growth rates 1-10 mm per year for massive slow 

growing corals

1-20 mm per year for Oculina and 

Lophelia3; growth rates of other taxa 

are unknown. 50-150 mm per year for faster 

growing branching corals

Number of reef building 

species

Approximately 800 Approximately 6-14

Nutrition Photosynthesis, zooplankton and 

suspended organic matter

Zooplankton and possibly suspended 

organic matter

Primary Threats1,2 Overfishing and destructive fishing Bottom-tending fishing gear

Pollution and siltation Oil and gas exploration and 

production

Coastal development Pipelines and cables 

Harvest of corals Climate change (ocean acidification 

and possible changes in currents and 

temperatures)

Recreational misuse

Diseases

Climate change (coral bleaching, 

ocean acidification and storm 

intensity)

1. Modified from Freiwald et al. 2004

2. 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. 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

corals, other deep coral taxa, such as stylasterids, 

gorgonians, and black corals do not form reefs, 

but often have complex, branching morphologies 

and may form dense groves or thickets. Sea 

fans may exist either singly on the seafloor or 

within large and complex ecosystems. The North 

Pacific, for example, is known to have extensive 

coral “gardens” composed of gorgonians and 

numerous other coral and sponge species. 

WHY ARE DEEP CORAL 

ECOSYSTEMS IMPORTANT?

As the understanding of deep coral 

communities and ecosystems has increased, 

so has appreciation of their value. Deep coral 

communities can be hot-spots of biodiversity 

in the deeper ocean, making them of particular 

conservation interest. Stony coral “reefs” as well 

as thickets of gorgonian corals, black corals, and 

hydrocorals are often associated with a large 

number of other species. Through quantitative 

surveys of the macroinvertebrate fauna, 

Reed (2002b) found over 20,000 individual 

invertebrates from more than 300 species living 

among the branches of ivory tree coral (Oculina 

varicosa) off the coast of Florida. Over 1,300 

species of invertebrates have been recorded in 

an ongoing census of numerous Lophelia reefs 

in the northeast Atlantic (Freiwald et al. 2004), 

and Mortensen and Fosså (2006) reported 361 

species in 24 samples from Lophelia reefs off 

Norway. Gorgonian corals in the northwest 

Atlantic have been shown to host more than 

100 species of invertebrates (Buhl-Mortensen 

and Mortensen 2005). An investigation by 

Richer de Forges et al. (2000) reported over 850 

macro- and megafaunal species associated with 

seamounts in the Tasman and south Coral Seas 

with many of these species associated with 

the deep coral Solenosmilia variabilis (Rogers 

2004). The three-dimensional structure of deep 

corals may function in very similar ways to 

their tropical counterparts, providing enhanced 

feeding opportunities for aggregating species, a 

hiding place from predators, a nursery area for 

juveniles, fish spawning aggregation sites, and 

attachment substrate for sedentary invertebrates 

(Fosså et al. 2002; Mortensen 2000; Reed 

2002b). 

The high biodiversity associated with deep coral 

communities is intrinsically valuable, and may 

provide numerous targets for chemical and 

biological research on marine organisms. For 

example, several deep-water sponges have 

been shown to contain bioactive compounds 

of pharmaceutical interest; sponges are often 

associated with deep coral communities. 

Bamboo corals (family Isididae) are being 

investigated for their medical potential as bone 

grafts and for the properties of their collagenlike gorgonin (Ehrlich et al. 2006). A number of 

deep corals are also of commercial importance, 

especially black corals (order Antipatharia) and 

pink and red corals (Corallium spp.), which are 

the basis of a large jewelry industry. Black coral 

is Hawaii’s “State Gem.”

Deep coral communities have also been 

identified as habitat for certain commerciallyimportant fishes. For example, commercially 

valuable species of rockfish, shrimp, and crabs 

are known to use coral branches for suspension 

feeding or protection from predators in Alaskan 

waters (Krieger and Wing 2002). Husebø et al. 

(2002) documented a higher abundance and 

larger size of commercially valuable redfish, ling, 

and tusk in Norwegian waters in coral habitats 

compared to non-coral habitats. Costello et al. 

(2005), working at several sites in the Northeast 

Atlantic, report that 92% of fish species, and 80% 

of individual fish were associated with Lophelia 

reef habitats rather than on the surrounding 

seabed. Koenig (2001) found a relationship 

between the abundance of economically 

valuable fish (e.g., grouper, snapper, sea bass, 

and amberjack) and the condition (dead, sparse 

and intact) of Oculina colonies. Oculina reefs 

off Florida have been identified as essential fish 

habitat for federally-managed species, as have 

gorgonian-dominated deep coral communities off 

Alaska and the West Coast of the United States. 

In other cases, however, the linkages between 

commercial fisheries species and deep corals 

remain unclear (Auster 2005; Tissot et al. 2006) 

and may be indirect.

Due to their worldwide distribution and the fact 

that some gorgonian and stony coral species 

can live for centuries, deep corals may serve as 

a proxy for reconstructing past changes in global 

climate and oceanographic conditions (Risk et 

al. 2002; Williams et al. 2007). The calcium 

carbonate skeletons of corals incorporate trace 

elements and isotopes that reflect the physical 

and chemical conditions in which they grew. 

Analysis of the coral’s microchemistry has 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

allowed researchers to reconstruct past oceanic 

conditions.

MAJOR GROUPS OF 

STRUCTURE-FORMING DEEP 

CORALS

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). Structureforming corals outlined in this document are 

animals in the cnidarian Classes Anthozoa 

and Hydrozoa that produce calcium carbonate 

(aragonite 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.

PHYLUM CNIDARIA

I. CLASS ANTHOZOA

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. 

I.A. SUBCLASS HEXACORALLIA

I.A.1. ORDER SCLERACTINIA 

 (STONY CORALS)

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 frameworkbuilding 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 

1995).

I.A.1.A. FAMILY CARYOPHYLLIIDAE

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 

1

Note 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 (Pourtalès, 1867).
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

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 (Fosså 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 56o S to 71o 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 “gracilis” 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 thicketlike 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 yr-1 (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 1.2 Samples of Lopehlia pertusa colonies collected from the Gulf of Mexico. The left 

specimen displays the more heavily calcified “brachycephala” morphology with large polyps, and 

the right specimen shows the more fragile “gracilis” morphology. Photo credit: Sandra Brooke, 

OIMB, Charleston, OR.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

grown continuously for the last 50,000 years 

(Schroder-Ritzrau et al. 2005). 

Reproduction: L. pertusa is a gonochoristic 

species (separate sexes) that produces a single 

cohort of about 3,000 relatively small (max = 

140 μm in diameter) oocytes per polyp each 

year (Waller and Tyler 2005). The species is an 

annual broadcast spawner, releasing gametes 

between January and February (Le Goff-Vitry 

and Rogers 2005; Waller and Tyler 2005). 

The low genetic diversity in some locations, 

the occurrence of genetically distinct fjord 

and offshore populations, and the presence 

of lecithotrophic larvae suggest there is a high 

degree of local recruitment (Le Goff-Vitry and 

Rogers 2005). Local recruitment, together 

with predominance of asexual reproduction via 

fragmentation, is thought to be critical in the 

persistence of populations, especially in areas 

impacted by trawling (Le Goff-Vitry and Rogers 

2005; Waller and Tyler 2005).

I.A.1.a.ii. Solenosmilia variabilis 

 Duncan, 1873

Description: Solenosmilia variabilis (Figure 

1.4) is a branching coral that often occurs as a 

secondary constituent of deep-water reefs. It is a 

prominent reef-building species on South Pacific 

seamounts.

Distribution: S. variabilis occurs throughout 

much of the Atlantic and Indo-Pacific Oceans, 

but is not found in the Arctic, Antarctic, and 

North and eastern Pacific waters (Cairns 1979). 

This coral forms dense clusters on Tasmanian 

Figure 1.3 Colonies of living and dead Lophelia with squat lobster. Photo credit: Ross et al., 

NOAA-OE.

Figure 1.4 Solenosmilia variabilis coral. Photo 

credit: Brooke et al., NOAA-OE, HBOI.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

 INTRODUCTION AND OVERVIEW

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

NOAA-OE, HBOI.

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 

available.

Reproduction: S. variabilis is a gonochoristic 

species with relatively small polyps (3.3 mm), 

small oocytes (148 μm), 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. FAMILY DENDROPHYLLIIDAE

I.A.1.b.i. Enallopsammia profunda 

 (Pourtalès, 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 deepwater 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 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

10

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

 (Pourtalès, 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 

8).

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 53o N (in the Atlantic) to 51o 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 

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 μm with an average 

of 144 oocytes per polyp.

I.A.1.C. FAMILY OCULINIDAE

I.A.1.c.i. Madrepora carolina 

 (Pourtalès, 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 coexisting 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 1.6 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.

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.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

11 INTRODUCTION AND OVERVIEW

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 69o N off 

Norway to 59o S latitude in the Drake Passage. 

Large individual colonies of M. oculata occur on 

exposed hard substrate throughout the Gulf of 

Mexico.

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 structurebuilding capability. 

Growth and Age: Limited information is 

available.

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 1.8a 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, 

HBOI.

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.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

12

(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: O. 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 deepwater 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 O. 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). 

Morphology: There are morphological 

differences between the shallow and 

deep-water colonies of O. 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 O. varicosa appears to 

be faster in deeper water (16 mm yr-1 at 

80 m) where zooxanthellae are absent, 

than at 6 m depth (11 mm yr-1). These 

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: O. 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

Figure 1.9 Oculina varicosa in the Oculina HAPC. Photo credit: 

L. Horn, NOAA Undersea Research Center at UNCWilmington.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

13 INTRODUCTION AND OVERVIEW

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. FAMILY POCILLOPORIDAE

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 structureforming 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 7o and 29o N latitude. In 

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 

1979). 

Morphology: M. myriaster is 

a branching species that forms 

broad, bushy colonies of 30-40 

cm in height (Cairns 1979). 

Growth, Age, and 

Reproduction: Limited 

information is available.

I.A.2. ORDER ZOANTHIDEA

Zoanthids are colonial, sea 

anemone-like anthozoans, 

mostly occurring in shallow 

tropical waters. While most of 

the more than 100 species of 

zoanthids do not form skeletal structures, deepwater gold corals are one taxon found in this 

order that does form rigid skeletons and grows 

to large sizes. 

I.A.2.A. FAMILY GERARDIIDAE

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 

Oceans.

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

Figure 1.11 Gold coral (Gerardia sp.) in Hawaii with a purple octocoral 

Clavularia grandiflora growing on it. Photo credit: A. Baco.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

14

been reported in the Hawaiian Archipelago and 

the Emperor Seamounts at depths of 350-600 m 

(Chapter 4) and in the Straits of Florida at depths 

around 600 m (Messing et al. 1990). 

Morphology: These corals form a rigid, 

branching, tree-like structure that can attain a 

height of up to 3 m (Figure 1.11).

Growth and Age: Gold corals appear to be very 

long-lived. A colony of Gerardia sp. collected off 

Little Bahama Bank was estimated to be 1,800 

years old (Druffel et al. 1995), and colonies off 

Hawaii have been aged at 450 to 2,742 years 

(Roark et al. 2006), placing them among the 

oldest known marine organisms.

Reproduction: Gold corals, like some other 

zoanthids, can be epizoic on other invertebrates; 

the larval stages settle out on other species 

of corals, particularly bamboo corals, and 

eventually overgrow the colony. Zoanthids are 

known to broadcast spawn during mass spawing 

events.Several species have separate sexes, 

while some are also hermaphroditic (Ryland and 

Babcock 1991). Reproductive strategies of gold 

coral are unknown.

I.A.3. ORDER ANTIPATHARIA 

 (BLACK CORALS)

Description: About 250 species of black coral 

are currently known. They do not form reefs, but 

like gorgonians and gold corals, large branching 

species can provide habitat for numerous other 

species. Though black corals are found in all 

U.S. regions they are best documented off 

Hawaii where they are commercially harvested 

for jewelry (Chapter 4; Grigg 2002). The order 

Antipatharia has recently been the subject of 

significant taxonomic revision (e.g., Opresko 

2001, 2002, 2003, 2004), and several new 

families have been proposed. A number of 

species in several families have been identified 

as important structure-forming corals in U.S. 

waters (Appendix 1.1 and 1.2).

Distribution: Antipatharians, commonly known 

as black corals, are found in all oceans, but are 

generally most common in deep-water habitats 

of tropical and subtropical oceans. They are 

generally anchored with a strong holdfast to hard 

substrate near drop-offs, terraces, ledges, and 

reef slopes in areas swept by strong currents. 

Black corals have recently been reported from 

Figure 1.12 A bushy black coral on Manning Seamount. Photo credit: The Mountains in the Sea Research 

Team, the IFE crew, and NOAA-OE.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

15 INTRODUCTION AND OVERVIEW

cruises to seamounts (Figure 1.12) in the Gulf 

of Alaska (Baco and Cairns 2005), Davidson 

Seamount off the California Coast (DeVogelaere 

et al. 2005), the New England Seamounts in the 

Atlantic (NOAA 2004), and in the northwestern 

Gulf of Mexico (E. Hickerson and G.P. Schmahl 

pers. comm.). 

Depth Range: Antipatharians are usually found 

at depths greater than 20 m, to a maximum of 

nearly 3,000 m (Etnoyer and Morgan 2005). 

Isolated colonies of deep-water species can 

be found in shaded areas as shallow as 4 m 

(Etnoyer and Morgan 2005), and a common 

temperate species from New Zealand (Antipathes 

fiordensis) is most abundant between 10 and 35 

m depth (Grange 1985).

Morphology: Antipatharians are hexacorals 

with branched (bushy, pinnate or fan-shaped) 

or unbranched (whip-like) skeletons covered 

with small spines or knobs and polyps that can 

be rust, yellow, green or white in color (Figure 

1.12). The polyps possess six unbranched, nonretractile tentacles, a feature that distinguishes 

them from gorgonians. The skeleton is black or 

dark brown in color and consists of chitin fibrils 

embedded in a protein matrix deposited as a 

series of layers or growth bands. 

Growth and Age: Black corals are known to 

achieve heights that exceed 3 m. Linear growth 

rates reported for black corals from temperate 

regions are much slower (e.g., Antipathes 

fiordensis; 1.6-3.0 cm yr-1; Grange 1997) than 

those of two commercially important species 

from Hawaii (Antipathes dichotoma, 6.42 cm 

yr-1 and Antipathes grandis, 6.12 cm yr-1; Grigg 

1976). While A. fiordensis is reported to reach 

sexual maturity at 70-105 cm, corresponding to 

a minimum age of 31 years (Parker et al. 1997), 

A. dichotoma and A. grandis are estimated to 

reach sexual maturity between 10-12.5 years 

(at heights of 64-80 cm), and can live about 

40 years (Grigg 1976). The age of another 

Hawaiian black coral occurring in deeper water 

(Leiopathes glaberrima) was recently estimated 

at around 2,377 years (Roark et al. 2006), and 

other species have been estimated to live longer 

than a century (Love et al. 2007; Williams et al. 

in press). 

Reproduction: A. fiordensis, a species from a 

New Zealand fjord, is a gonochoristic broadcast 

spawner with seasonal reproductive patterns 

(Parker et al. 1997). Mature oocytes are 100-

140 μm in diameter and female colonies 

produced 1.3-16.9 million oocytes. As with all 

colonial corals, the larger colonies dominate 

the reproductive output of the population (Miller 

1996). 

I.B. SUBCLASS OCTOCORALLIA

The subclass Octocorallia includes gorgonians 

(sea fans and sea whips), true soft corals, 

stoloniferans, and sea pens, all groups that 

include deep-water species, as well as the order 

Helioporacea. The latter includes the small family 

Lithotelestidae (Bayer and Muzik 1977) with at 

least one deep coral species in the Caribbean 

(Chapter 8) and the family Helioporidae, which 

contains one extant shallow-water species, the 

blue coral. Octocorals are distinguished from 

other anthozoans by the presence of eight featherlike (pinnate) tentacles. Octocorals can form 

large, long-lived colonies with many (thousands) 

of tiny polyps, but they do not form complex 

reef structures. They all contain calcareous 

spicules within their tissue (coenochyme) and 

some (order Gorgonacea) also have a central 

proteinaceous rod with embedded calcareous 

spicules or heavily calcified skeletal elements 

that alternate with non-calcified gorgonin 

elements. Currently about 2,700 species of 

octocorals have been described; most occur in 

shallow water, although several hundred species 

also occur in deep water. In deep-water habitats 

where stony corals are less abundant, such as 

seamounts and at high latitudes, octocorals 

are more prevalent and form the basis of the 

coral ecosystem. Gorgonians, true soft corals, 

and stoloniferans are now commonly grouped 

together within the single order Alcyonacea 

(Bayer 1981; Fabricius and Alderslade 2001), but 

are discussed as separate taxa in this report to 

provide additional detail regarding the distribution 

of these important functional groups of corals. 

The orders Helioporacea and Pennatulacea are 

clearly delineated as separate orders from the 

remaining octocorals.

I.B.1. ORDER ALCYONACEA 

 (TRUE SOFT CORALS)

In general, these soft corals are less important 

structure-forming species than are many 

gorgonians, although the families Alcyoniidae 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

16

and Nephtheidae include deep-water species 

that achieve relatively large sizes (Watling 

and Auster 2005). Soft corals of the genus 

Eunephthea (formerly Gersemia) are widespread 

and are the most abundant corals in the Bering 

Sea (Chapter 2). True soft corals of the order 

Alcyonacea generally lack a rigid internal skeleton 

for support, but have separate calcareous 

spicules embedded in the fleshy coenochyme. 

Stoloniferans, now included in the Alcyonacea, 

have small polyps that are often connected to 

each other by a thin runner or stolon. With the 

exception of the tropical shallow-water organpipe coral (Tubipora musica), most are not 

important structure-forming corals. However, 

a few species can form extensive mats on 

hard surfaces such as rocks, other corals, and 

sponges (Stone 2006).

I.B.2. ORDER GORGONACEA 

 (SEA FANS)

Major structure-forming families in the order 

Gorgonacea include Isididae, Coralliidae, 

Paragorgiidae, and Primnoidae (Morgan et al. 

2006), with species in the families Plexauridae, 

Acanthogorgiidae, Ellisellidae, Chrysogorgiidae, 

and Anthothelidae providing structure to some 

degree (Appendix 1.1 and 1.2). At least 12 

families are known to occur in waters deeper 

than 200 m (Etnoyer et al. 2006). Gorgonians 

are the most important structure-forming corals 

in the Gulf of Alaska and the Aleutian Islands, 

where they form both single- and multi-species 

assemblages (Chapter 2). 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). Most gorgonians 

have a solid proteinaceous (gorgonin) central 

axis with embedded calcareous sclerites that 

provide support, covered by a thin layer of tissue 

(coenenchyme and polyps) with embedded 

calcareous spicules (Fabricius and Alderslade 

2001). They often exhibit a branching 

morphology, can occur at high density and cover, 

and reach considerable size (>3 m tall), thus 

providing structure and habitat for associated 

fauna.

I.B.2.A. FAMILY ISIDIDAE 

 (BAMBOO CORALS) 

Description: Isididae is a large family with over 

150 species of mostly deep-water corals. The 

most common deep-water genera are Acanella, 

Isidella and Keratoisis. Acanella arbuscula, a 

species occurring in the northwestern Atlantic 

(Chapters 5 and 7) is unusual in that it anchors 

in mud rather than on hard substrata (Mortensen 

and Buhl-Mortensen 2005a). Several species 

are collected for jewelry.

Distribution: Bamboo corals are thought to 

have a cosmopolitan distribution and important 

structure-forming species have been identified 

in the Gulf of Mexico, the Southeast, Hawaii, 

the West Coast, the northeast Pacific and IndoPacific (Fabricius and Alderslade 2001; Etnoyer 

and Morgan 2003; Appendix 1.1 and 1.2). 

Depth Range: In general bamboo corals occur 

below 800 m (Etnoyer and Morgan 2005), with 

the deepest recorded at 4,851 m (Bayer and 

Stefani 1987). In Alaska bamboo corals are 

observed between 400 and 2,827 m but have 

been collected from depths of 3,532 m (Chapter 

2). However, four genera have been reported 

from tropical Indo-Pacific reefs at depths of 10-

120 m (Fabricius and Alderslade 2001).

Morphology: Colonies can be whip-like but are 

usually branched, bushy or fan-like (Figure 1.13) 

and can range in size from tens of centimeters 

to over a meter (Verrill 1883). Colonies have a 

distinctly articulated skeleton of heavily calcified 

internodes and proteinaceous gorgonin nodes 

(a stiff leathery matrix consisting of protein and 

mucopolysaccharides). The alternating segments 

give the isidid branches a unique bamboo-like 

appearance (Figure 1.13), hence the name 

“bamboo-coral.”

Growth and Age: Recent studies by Andrews 

et al. (2005a,b) have estimated radial growth 

rates for bamboo corals that ranged from 

approximately 0.05 (age 150 years) to 0.117 mm 

yr-1 (age 43 years). Linear growth rates up to 30 

mm yr-1 have been estimated for Lepidisis sp. in 

New Zealand waters (Tracey et al. in press).

Reproduction: Reproductive strategy is thought 

to be similar to that of other octocorals with 

colonies having separate sexes and gametes 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

17 INTRODUCTION AND OVERVIEW

being broadcast into the water column in a 

synchronous manner (Fabricus andAlderslade 

2001). 

I.B.2.B. FAMILY CORALLIIDAE 

 (RED AND PINK CORALS)

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 19o 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 

A

B

Figure 1.13 A. Bamboo coral (Keratoisis

sp.) on the Davidson Seamount (1,455 

meters). Coral colony age estimates exceed 200 years. Image courtesy of NOAA, 

MBARI 2002. B. Close-up view of white 

bamboo coral (Keratoisis flexibilis) showing 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.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

18

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 

yr-1 (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. 

B

Figure 1.14a Pink coral 

necklaces for sale in 

Japan. Photo credit: Andy 

Bruckner, NOAA.

Figure 1.14b Corallium sp., with deep 

purple Trachythela

octocoral, brittle 

stars, crinoids and 

sponges. Photo 

credit: The Mountains in the Sea 

Research Team, 

the IFE crew, and 

NOAA-OE.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

19 INTRODUCTION AND OVERVIEW

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).

I.B.2.C. FAMILY PARAGORGIIDAE 

 (BUBBLEGUM CORALS)

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 

of 200-1,100 m, where it can occur in dense 

2

In earlier studies, more than 50% of colonies were reported 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 reproductive maturity by 3-4 years (Marschal et al. 2004). 

Figure 1.15 a. Paragorgia, or “bubblegum corals,” grow to 

over 2.5 m tall. On Davidson Seamount, where this photo 

was taken, they are found primarily on the highest 

elevations. b. Close up of Paragorgia. Photo credit: The 

Davidson Seamount Expedition, MBARI, and NOAA-OE.

A

B
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

20

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. 

2005). 

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 BuhlMortensen (2005b) report estimates of linear 

growth rates for P. arborea in New Zealand 

and Norway between 2.2-4.0 cm yr-1 and 0.8-

1.3 cm yr-1 respectively. Andrews et al. (2005a) 

estimated a Pacific species to have grown 

at a minimum of 0.5 cm yr-1 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 

2001).

I.B.2.D. FAMILY PRIMNOIDAE 

Description: Primnoidae is a large family (>200 

species) that includes a number of conspicuous 

and abundant branching species in the genera 

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.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

21 INTRODUCTION AND OVERVIEW

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 

2001).

I.B.3. ORDER PENNATULACEA 

 (SEA PENS)

Description: Pennatulaceans are a diverse but 

poorly known group of octocorals that include 

16 families, most of which live in the deep sea. 

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 

worldwide.

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 

Service.

3

The authors identified this species in Alaska as Primnoa 

resedaeformis, which is now thought to occur only in the 

Atlantic.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

22

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 stemlike base or foot (peduncle) 

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 

yr-1 depending on size with an 

estimated longevity approaching 

50 years for moderately sized 

specimens.

Reproduction: Sea pens are known to be 

broadcast spawners but other reproductive 

information is limited. 

II. CLASS HYDROZOA

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). 

II.A.1. ORDER ANTHOATHECATAE 

II.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 

Figure 1.18 Stylaster coral. Photo credit: Phillip Colla Photography, 

Oceanlight, Carlsbad, CA.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

23 INTRODUCTION AND OVERVIEW

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 

superficial.

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.

comm.).

Growth and Age: Limited information is 

available. 

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).

THREATS TO DEEP CORAL 

ECOSYSTEMS

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 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

24

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.

Major types 

of bottomtending 

fishing gear

Gear types that 

may impact 

deep corals Description of gear

Description of impact on 

corals

Potential 

severity 

of disturbance

Trawls

Bottom/ 

otter trawls

A large net is held open by two doors 

and dragged behind a fishing boat 

along the seafloor; gear in contact 

with seafloor can be 30-100 m in 

length; primarily used to harvest 

demersal finfish and rock shrimp

Incidentally removes, displaces or damages 

corals**

High

Mid-water trawls

Similar type of gear to the bottom 

trawl but designed to harvest pelagic 

fish species; no protective gear on 

the footrope

May come in contact with 

bottom during fishing, 

which can remove, displace or damage corals §

Low

Dredges Scallop dredges

A large steel frame is dragged behind 

a fishing boat along the seafloor; 

specifically used to harvest scallops

Incidentally removes, 

displaces or damages 

corals S 

High

Longlines

Bottom-set longlines/ 

demersal longlines

A nylon or poly line with up to 

thousands of attached hooks; deployed along the seafloor in lengths 

up to 2-5 km

May entangle or detach 

corals during retrieval † Low

Traps or 

pots

Single-set pots

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

Limited spatial damage to 

corals during pot retrieval 

‡

Low

Longline pots Single pots are strung together on a 

long line (10-90 pots)

Damages corals during 

pot retrieval and entanglement with lines; under 

certain conditions gear 

can be dragged like a 

plough on seafloor §

Medium

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; † High 1998; § NMFS 2004; ‡ Eno et al. 2001; 

S Stone et al. 2005.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

25 INTRODUCTION AND OVERVIEW

Bottom Trawls and Dredges

The National Research Council (2002) concluded 

that bottom trawling and dredging reduce habitat 

complexity by removing or damaging the actual 

physical structure of the seafloor, and it causes 

changes in species composition. Stable 

communities of sessile, long-lived species, 

such as corals are especially vulnerable to 

acute and chronic physical disturbance (NRC 

2002). Mobile bottom-tending fishing gear 

(trawling/dredging) reduces habitat complexity 

by removing structure-forming organisms that 

provide shelter for fishes and invertebrates 

(Figure 1.19). Areas of the seafloor with rough 

or steep bathymetry or composed mostly of 

bedrock and boulders are infrequently trawled 

due to the risk of damaged and lost gear. Such 

areas may support coral habitat and serve as 

de facto reserves.

Recovery of a trawled or dredged area, if 

allowed to occur, can take a few months to 

several decades, if it occurs at all, depending 

on the intensity and frequency of disturbances 

to the seafloor (Hutchings 2000). In general, 

there have been limited studies on the longterm impacts of trawling and dredging (FAO 

2005). Recovery rates range from one to five 

times their generation time, depending on the 

life history of a particular organism (Emeis et 

al. 2001). In more complex habitats such as 

deep coral communities full recovery may require 

decades to centuries due to their slow growth 

rates (Freiwald et al. 2004). 

Studies from around the world have reported 

severe disturbance to deep coral communities 

from trawling (ICES 2005). Disturbance 

from bottom trawling has been documented 

on Solenosmilia coral habitats on Australian 

seamounts (Koslow et al. 2000, 2001; Anderson 

and Clark 2003); gorgonian forests in New 

Zealand (Probert et al. 1997); off Alaska’s Aleutian 

Islands, Alaskan primnoid habitats (Krieger 

1998, 2001), corals in boulder habitats off Alaska 

(Freese et al. 1999), and coral gardens off Alaska 

(Stone 2006); Oculina reefs off Florida (Koenig 

et al. 2005); Lophelia reefs in the northeast 

Atlantic (Rogers 1999; Hall-Spencer et al. 2001; 

Fosså et al. 2002; Grehan et al. 2005; Wheeler 

et al. 2005) and Atlantic Canadian waters (Butler 

and Gass 2001). The distribution of Lophelia

reefs has likely been reduced by trawling on the 

European continental slope (Roberts et al. 2003; 

ICES 2005).

Traps and pots

Disturbance to deep coral communities from 

single pot fishing is expected to be much less 

Figure 1.20 Steel pots are used to harvest many species of crabs in Alaska. Some pots, such as this one, 

measure 2 x 2 x 1 m and may weigh more than 300 kg. 

This pot was derelict for some time and has been heavily 

colonized by soft corals. Photo credit: Alaska Department of Fish and Game

Figure 1.19 Red tree corals (Primnoa sp.) are periodically caught with trawl gear in Alaskan waters. 

This specimen was caught during a NOAA Fisheries 

groundfish stock assessment survey in Dixon 

Entrance, Gulf of Alaska. Photo credit: R. Lauth, 

Alaska Fisheries Science Center.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

26

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).

Harvest

Certain deep corals have been collected or 

harvested for jewelry and curios since antiquity. 

In the U.S., commercial harvest of precious 

Figure 1.21 The black coral divers of Lahaina, Maui. 

Team leader Robin Lee is the diver wearing the cap. 

Photo credit: R. Grigg. 

4

Precious corals refer to red and pink corals (family Coralliidae). The term is often used more broadly to also include 

black corals (family Antipathidae), gold corals (family Gerardiidae), and bamboo corals (family Isididae).

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 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

27 INTRODUCTION AND OVERVIEW

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 

Figure 1.22 Map of the Gulf of Mexico showing active leases by water depth in 2005. Image credit: Minerals 

Management Service.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

28

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 hardsubstrates. 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 cobaltenriched 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).

At least two companies have plans to begin 

exploration or mining in the territorial seas of 

other countries in the near future (Schrope 

2007), but there are no current plans for mining 

within the U.S. EEZ. Any exploitation of mineral 

resources outside areas of national jurisdiction 

would be governed by guidelines established by 

the International Seabed Authority (ISA) (www.

isa.org.jm). The Authority is an autonomous 

international organization established under the 

1982 United Nations Convention on the Law of 

the Sea (UNCLOS) and the 1994 Agreement 

relating to the Implementation of Part XI of 

UNCLOS. Through the ISA Parties to the 

Convention administer 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).

5

More information on this program is available online at 

http://www.mms.gov/sandandgravel/
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

29 INTRODUCTION AND OVERVIEW

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 

introduction of non-indigenous species should 

be the priority. 

The invasive snowflake coral (Carijoa riisei; 

Figure 1.23) was first discovered in Pearl Harbor 

in 1972 on pier pilings and is believed to have 

arrived from the Indo-Pacific area as a hull 

fouling organism. The snowflake coral most 

often occurs in shaded areas, and has rapidly 

spread to deeper waters where it settles on and 

eventually smothers black coral colonies. In a 

survey of the Maui Black Coral Bed in 2001, up 

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 

semi-mobile epifauna (Figure 1.24). 

In large parts of the affected area, the 

tunicate covers 50% or more of the 

substrate. Didemnum sp. may be a 

serious threat to deep coral that occur 

on hard substrates in the Northeast, 

particularly for those corals (such as 

Paragorgia and Primnoa) that are 

known to occur on the gravel substrate 

of the Northeast Peak of Georges Bank. 

It has the potential to spread rapidly by 

budding and fragmentation of the mats 

could promote rapid range expansion.

The rates of invasions and vectors of 

transmission for invasive species in 

the deep sea are unknown. Because 

limited knowledge exists about the 

biology and ecology of deep corals in 

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 0.1°C 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, 

U.S. Geological Survey.

Figure 1.23 Black coral at approximately 100 m depth overgrown with the invasive snowflake coral Carijoa riisei. Photo 

credit: HURL (Hawaii Undersea Research Laboratory) archives, 

R. Grigg and S. Kahng.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

30

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 (meridional 

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 

corals. 

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 largescale 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 “aragonite 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 carbonatebased 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 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

31 INTRODUCTION AND OVERVIEW

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 

disturbances. 

National Overview

In the chapters that follow, the authors draw 

together current knowledge, including previously 

unpublished data, on the distribution of deep 

Figure 1.25 Map showing the U.S. EEZ and areas of seven regional assessments.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

32

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, their associated 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.

THE DISTRIBUTION OF DEEP CORAL 

COMMUNITIES IN U.S. WATERS

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.

Pacific

The major oceanographic influences on the 

U.S. Pacific Coast are the Alaska and California 

Currents, which are formed when the eastwardflowing 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 

biomass. 

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 reefbuilders had been reported. Stony corals accrete 

calcium carbonate in the form of aragonite, and 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

33 INTRODUCTION AND OVERVIEW

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 

there.

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 nonemergent 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 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

34

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 deepwater 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. 

Atlantic

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. 

Figure 1.26 Paragorgia sp. crown a ridge on the Davidson Seamount. Photo credit: The Davidson Seamount 

Expedition, MBARI, and NOAA-OE.

6

For more information on Large Marine Ecosystems and 

their designation visit www.edc.uri.edu/lme
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

35 INTRODUCTION AND OVERVIEW

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 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

36

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 Pourtalès 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 1.28 Multi-colored gorgonians with whip coral in background. East Bank, Flower Garden Banks NMS

Photo credit: NURC/UNCW and NOAA/FGBNMS 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

37 INTRODUCTION AND OVERVIEW

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.

U.S. CONSERVATION AND

MANAGEMENT MEASURES

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 

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 deepwater fishes. Photo credit: FGBNMS and NURC/UNCW.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

38

the world (e.g., Rogers 1999; Koslow et al. 2000; 

Hall-Spencer et al. 2001; Fosså et al. 2002; 

Roberts 2002; Grehan et al. 2005, Wheeler et al. 

2005). Harvest of black and precious corals in 

Hawaii has been identified as a moderate threat, 

but it is conducted in a very selective manner, 

and its overall impact is minor compared to trawl 

fishing in other regions. Hawaii is also the only 

jurisdiction that has specifically identified a current 

threat to deep corals from an invasive species. 

Oil and gas exploration and development in the 

Gulf of Mexico, where it is increasingly moving 

into deeper waters, is the only non-fishing direct 

anthropogenic stressor that poses a moderate 

threat to deep coral communities. Potential 

impacts from climate change (Including ocean 

acidification) are largely unknown.

U.S. Management of Deep Coral Ecosystems

in an International Context 

Recent interest in deep coral ecosystems has 

galvanized the public and triggered conservation 

and management action in the United States and 

around the world. In recent years, conservation 

actions have been taken shortly after discovery 

of vulnerable deep coral habitats. Internationally, 

this includes new marine protected areas 

established to protect deep coral communities 

in the northeast and northwest Atlantic, in the 

Canadian Pacific, and on seamounts in Australia 

and New Zealand. Internationally, a series 

of United Nations General Assembly (UNGA) 

resolutions has addressed the impacts of fishing 

on vulnerable marine ecosystems in international 

waters, with specific reference to seamounts, 

hydrothermal vents, and cold-water corals. This 

international effort culminated in the December 

2006 UNGA Sustainable Fisheries Resolution 

(A/61/105), which calls upon states and regional 

fisheries management organizations (RFMOs) 

to ensure the sustainable management of fish 

stocks and protection of vulnerable marine 

ecosystems – including seamounts, hydrothermal 

vents, and cold-water corals – from destructive 

fishing practices by December 31, 2008. 

In the United States, on October 6, 2006, 

President Bush put forth a memorandum to 

promote sustainable fishing and end destructive 

Regions

Threats Alaska

West 

Coast

Pacific 

Islands Northeast Southeast

Gulf of 

Mexico Caribbean

Bottom trawl 

fishing impacts High High NA High High

Low - 

Medium NA

Other bottom 

fishing impacts

Low - 

Medium

Low - 

Medium Low

Low - 

Medium

Low - 

Medium

Low - 

Medium Low

Deep coral harvest NA NA Medium NA NA NA NA

Oil and gas 

development Low Low NA NA NA Medium NA

Cable deployment Low Low Unknown Low Low Low Unknown

Sand and gravel 

mining Low NA NA Low Low Low NA

Invasive species Unknown Unknown Medium Unknown Unknown Unknown Unknown

Climate change Unknown Unknown Unknown Unknown Unknown Unknown Unknown

Table 1.3. Summary of perceived levels of current threats to deep coral communities for U.S. regions. NA = Not 

Applicable (i.e., this threat is prohibited or does not occur anywhere within that region). Threat levels are based 

on the information provided by the regional chapter authors. 

Note: These perceived threat levels reflect only the occurrence of these stressors in a region, and their potential, 

if unmitigated, to damage deep coral communities they might encounter. They do not indicate the actual impacts 

of each stressor, which will likely vary widely within and among regions. Since the location of deep coral habitats 

is incompletely known, there is uncertainty over their degree of overlap with human activities. Substantial 

management steps have been taken to mitigate threats. For example, significant actions to minimize adverse 

impacts of bottom fishing gear through gear modifications and gear closures have been taken in each region, 

and management procedures are in place to mitigate potential impacts of oil and gas development and mining 

where they occur on the outer continental shelf.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

39 INTRODUCTION AND OVERVIEW

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 

(P.L. 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 O. 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 

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 (P.L. 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 deepsea 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, 

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.

nmfs.noaa.gov/pr/species/concern/#corals).
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

40

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 deepsea 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 

areas. 

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 

Regional Fishery 

Management 

Council

Deep Coral or 

Biogenic Habitat 

Identified as EFH for 

Managed Species?

Coral 

Management 

Plan?

Extensive Areas Protected 

from Gear Impacts?

Coral Bycatch 

Reasonably Well 

Monitored

North Pacific Yes - Groundfish No Yes Yes

Pacific Yes - Groundfish No Yes Yes

Western Pacific No Yes – Precious 

Corals Yes Not Applicable

New England No No

Limited – Lydonia & 

Oceanographer Canyons 

and New England Groundfish 

Habitat Closure Areas

No

Mid Atlantic No No No No

South Atlantic Yes- Snapper 

Grouper Complex

Yes - Used to 

protect Oculina 

Banks

Limited - Oculina Banks 

HAPC No

Gulf of Mexico No

Yes - Not yet 

applied to deep 

coral

No No

Caribbean No

Yes - Not yet 

applied to deep 

corals

No Not Applicable

Table 1.4. Regional fishery management council management actions affecting deep coral habitat.
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

41

found in the Papahānaumokuākea Marine 

INTRODUCTION AND OVERVIEW

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 

Figure 1.30 Rockfish take refuge among a primnoid octocoral in Olympic Coast National 

Marine Sanctuary. Photo credt: OCNMS/NOAA
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

42

Fishery Management Councils in the Pacific to 

propose historic protective measures limiting 

bottom-trawling in areas that might contain coral 

resources. 

The U.S. West Coast and Alaska have an 

extensive history of fisheries using bottomcontact gear, including bottom trawling for Pacific 

cod, hake and rockfishes; bottom-set longlines 

for fish; and individual traps and multiple traplines for crab in Alaska. Alaska is currently home 

port to the largest fleet of U.S. bottom trawlers. 

The importance of these bottom-trawl fisheries 

has been a major factor in the development 

of NMFS trawl surveys. These surveys have 

provided the broadest scale information on the 

distribution and abundance of deep corals in 

these two regions (Chapters 2 and 3).

The North Pacific Fishery Management Council 

has taken a number of important steps that reduce 

the impact of fisheries on essential fish habitat in 

the EEZ around Alaska. Beginning in 1998, the 

Council prohibited trawling in the eastern Gulf of 

Alaska and southeast Alaskan waters within a 

180,400 km2 area as part of a license-limitation 

program. The measure was originally proposed 

in 1991 over conservation concerns for rockfish 

stocks to protect seafloor habitat from long-term 

disturbance from trawling. In 2000 the Council 

established the 10.6 km2 Sitka Pinnacles Marine 

Reserve in the Gulf of Alaska and prohibited all 

bottom-fish gear types (except pelagic troll gear 

for salmon) in the reserve. These pinnacles 

consist of two large volcanic cones that rise to 

within 40 and 70 m of the ocean surface, and 

provide a variety of high-relief habitats colonized 

by the deep coral Primnoa sp., anemones, and 

other organisms. Aggregations of lingcod and 

several juvenile and adult rockfish species are 

associated with the pinnacles. 

Recently, the North Pacific and Pacific Fishery 

Management Councils each took historic steps, 

recommending to “freeze the footprint” of bottomtrawling within their respective jurisdictions in 

order to protect EFH. In 2006, NMFS approved 

a number of North Pacific Council recommended 

EFH closures in the Aleutian Islands and Gulf 

of Alaska. Many of these areas are known 

to contain important deep coral and sponge 

habitats. More than 950,000 km2 along the 

remote Aleutian Islands were closed to bottom 

trawling – targeting areas that had not yet 

received extensive trawling, with 377 km2 of “coral 

gardens” closed to all bottom-tending fishing 

gear. Additionally, 7,155 km2 in Gulf of Alaska 

Slope Habitat Conservation Areas were closed 

to bottom trawling and 18,278 km2 of Alaska 

seamounts and 46 km2 of Primnoa coral areas 

in the Gulf of Alaska were closed to all bottomtending fishing gear. In June 2007, the North 

Pacific Fishery Management Council adopted 

additional measures to conserve benthic fish 

habitat in the Bering Sea. These measures, if 

approved, NMFS would prohibit bottom trawling 

over an additional area of more than 450,000 

km2.

The Pacific Fishery Management Council is 

responsible for developing FMPs for fisheries off 

the coasts of California, Oregon, and Washington. 

Within the past three to six years, commercial 

fishing has been prohibited or significantly 

curtailed within the Cowcod and Rockfish 

Conservation Areas. While these restrictions 

were not designed to address impacts on deep 

corals, they are likely to protect some deep coral 

habitats. Beginning in 2000, the Council also 

prohibited footrope trawls (footrope=weighted 

edge of trawl that impacts seafloor) greater 

than 8 inches on most of the continental shelf, 

effectively making many complex, rocky habitats 

that are home to deep corals inaccessible to 

trawlers. 

In 2006, NMFS approved a plan that identified 

and described EFH for Pacific groundfish and 

prohibited bottom trawling in 336,700 km2 of 

habitat off the West Coast of the U.S. This 

represents over 42 percent of the EEZ off 

Washington, Oregon, and California, including 

areas that may contain deep coral and sponge 

habitats. Selected areas with known deep 

coral resources (e.g., Davidson Seamount) are 

protected from all bottom-contact gear. 

Unlike most other areas of the United States, the 

Insular Pacific has no history of domestic bottomtrawl fisheries. The Western Pacific Fishery 

Management Council manages the fisheries in 

federal waters around the Territory of American 

Samoa, Territory of Guam, State of Hawaii, 

Commonwealth of the Northern Mariana Islands, 

and other U.S. Pacific island possessions. It has 

the oldest and most comprehensive restrictions 

designed to protect coral and other biogenic 

habitat from adverse impacts of fishing gear. 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

43 INTRODUCTION AND OVERVIEW

In 1983, the Council prohibited the use of trawl 

gear, bottom-set longlines, and bottom-set 

gillnets - all identified as threats to deep corals 

- within all waters in their region of the U.S. EEZ. 

This action was taken, in part, in response to 

observed impacts of foreign trawl fisheries on 

seamounts (e.g., Hancock Seamount) before the 

declaration of the U.S. EEZ. 

In 1983, the Western Pacific Fishery Management 

Council also developed a Precious Corals FMP. 

The coral beds included in the FMP contain 

several deep coral species (Chapter 4). Under 

the 1983 FMP and its amendments, NMFS 

established quotas and minimum legal sizes for 

harvest of pink, black, gold, and bamboo coral, 

and, in 2002, prohibited the use of non-selective 

gear. Currently, only the black coral fishery 

within Hawaii State waters is active, and the 

Council and State are in the process of revising 

management plans that incorporate the recent 

impacts of the invasive soft coral Carijoa riisei.

Regional Management Actions 

in the U.S. Atlantic

The Northeast U.S. region has the longest history 

of major trawl and scallop dredge fisheries in 

the United States and its bottom-trawl fishery 

is second in size only to Alaska’s. As a result, 

much of the continental shelf has been heavily 

trawled or dredged. In addition to trawling, there 

are active fisheries using bottom-set longlines, 

gillnets, and pots and traps, some extending into 

the slope and canyon habitats that are known 

deep coral habitat. 

The New England Fishery Management Council 

manages fisheries off the coast of Maine, New 

Hampshire, Massachusetts, Rhode Island, 

and Connecticut. The Mid-Atlantic Fishery 

Management Council is responsible for 

management of fisheries in federal waters off the 

coasts of New York, New Jersey, Pennsylvania, 

Delaware, Maryland, Virginia, and North 

Carolina. 

The two Councils oversee significant trawl 

and dredge fisheries with potential impacts 

on deep coral habitats. In 2005, in order to 

minimize adverse impacts to EFH, NMFS 

approved Council-recommended closures 

of Oceanographer and Lydonia Canyons 

(approximately 400 km2, on the southern flank of 

Georges Bank), to bottom trawling and gillnetting 

for monkfish. These canyons are areas of known 

deep coral communities. Also approved were 

limits on the size of the bottom-trawling roller 

gear and rockhopper gear on the footrope of the 

nets to no more than six inches in diameter in 

the submarine canyon areas off the shores of 

the mid-Atlantic states known as the “southern 

management area” of the monkfish fishery. In 

The South Atlantic Fishery Management Council 

has jurisdiction over FMPs in the EEZ off North 

Carolina, South Carolina, Georgia, and eastern 

Florida to Key West (note: North Carolina is 

represented in both the Mid-Atlantic and South 

Atlantic Councils). Trawling in the south Atlantic 

region is primarily limited to shrimp trawling 

on the continental shelf. The Council was an 

early leader in addressing the threats of bottom 

trawling to deep coral communities. In 1984, the 

Council established the 315 km2 Oculina HAPC, 

the world’s first protection granted specifically to 

a deep coral habitat. In 2000 the South Atlantic 

Council expanded the Oculina HAPC to 1,043 

km2. The South Atlantic Council is currently 

reviewing and evaluating options for gear 

regulations and four new HAPCs containing deep 

coral habitats, including two very large areas, 

as part of a Comprehensive Fishery Ecosystem 

Plan Amendment (Chapter 6).

The relatively shallow nature of the Oculina coral 

banks probably led to their early recognition as 

important habitats to conserve. Unfortunately, 

they have also been more accessible to trawlers 

(primarily rock shrimp and calico scallop). Despite 

early protection, enforcement difficulties resulted 

in continued destruction through illegal fisheries 

until recent requirements for use of vessel 

monitoring systems and enhanced enforcement. 

Koenig et al. (2005) estimated that 90% of 

Oculina coral banks had been damaged by 

trawling by 2001 and only 10% remained intact. 

This is perhaps the clearest U.S. example of the 

extensive damage to deep coral communities 

by trawling. Oculina varicosa was identified 

by NMFS in 1991 as a “candidate species” for 

potential listing under the Endangered Species 

Act, based on well-documented declines in the 

Oculina coral banks areas due to damage from 

mechanical fishing gear, coupled with a lack of 

observed recruitment. In 2000 this designation 

was revised to “species of concern.”
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

44

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 shallowwater 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. 

DEEP CORAL INFORMATION NEEDS AND 

RESEARCH PRIORITIES

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 

technology. 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

45 INTRODUCTION AND OVERVIEW

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, 

Figure 1.31 3-D colored bathymetry of Chapman’s Reef, from 2005 survey done with multibeam 

sonar from R/V Cape Fear by Seafloor Systems, Inc. Image credit: A. Maness. 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

46

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 groundtruthing 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 

techniques.

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 webaccessible 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

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 

absent. 

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 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

47 INTRODUCTION AND OVERVIEW

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 followon 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 sourcesink 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 

corals to disturbance. These studies include 

factors influencing reproduction, recruitment, 

and recolonization rates, as well as patterns and 

processes of growth and mortality for key coral 

species.

Biodiversity and Ecology of Deep Coral 

Communities

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 MagnusonStevens 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 

Acidification 

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 longlived gorgonians and over geologic timescales in 

stony coral reef mounds (Smith et al. 1999; Risk 

et al. 2002; Williams et al. 2006).
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

48

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 longstanding 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 

areas where deep coral communities are not 

threatened.

REFERENCES

Adkins JF, Henderson GM, Wang S, O’Shea S, 

Mokadem F (2004) Growth rates of the 

deep-sea Scleractinia Desmophyllum 

cristagalli and Enallopsammia rostrata. 

Earth and Planetary Science Letters 

227:481–490

Alcock A (1902) Report on the deep-sea 

Madreporia of the Siboga Expedition. 

Siboga-Expeditie 16a:1–52

Andrews AH, Cordes EE, Mahoney MM, Munk K, 

Coale KH, Cailliet GM, Heifetz J (2002) 

Age, growth and radiometric age validation 

of a deep-sea, habitat-forming gorgonian 

(Primnoa resedaeformis) from the Gulf of 

Alaska. Hydrobiologia 471:101–110

Andrews, AH, Cailliet GM, Kerr LA, Coale KH, 

Lundstrom C, DeVogleare A (2005a) 

Investigations of age and growth for 

three species of deep-sea coral from the 

Davidson Seamount off central California. 

Pages 965–982 in Freiwald A, Roberts JM 

(eds.), Cold-water corals and ecosystems. 

Springer-Verlag Berlin Heidelberg

Andrews AH, Tracey DM, Neil H, Cailliet GM, 

Brooks CM (2005b) Lead-210 dating 

bamboo coral (family Isididae) of New 

Zealand and California (abstract) Third 

International Symposium on DeepSea Corals: Science and Management. 

University of Miami, Florida

Anderson OF, Clark MR (2003) Analysis of 

bycatch in the fishery for orange roughy, 

Hoplostethus atlanticus, on the South 

Tasman Rise. Marine and Freshwater 

Research 54(5):643–652

Auster PJ (2005) Are deep-water corals important 

habitats for fishes? Pages 747–760 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin Heidelberg
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

49 INTRODUCTION AND OVERVIEW

Auster PJ, Moore J, Heinonen KB, Watling L 

(2005) A habitat classification scheme 

for seamount landscapes: assessing the 

functional role of deep-water corals as 

fish habitat. Pages 761–769 in Freiwald 

A, Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Avent R, King M, Gore R (1977) Topographic and 

faunal studies of shelf-edge prominences 

off the central eastern Florida coast. 

Internationale Revue der Gesamten 

Hydrobiologie 62:185–208

Baco A, Cairns SD (2005) Distribution of corals on 

Derickson Seamount, a deep seamount 

near the Aleutian chain of Alaska (abstract) 

Third International Symposium on Deepsea Corals: Science and Management. 

University of Miami, Florida

Baco A, Shank TM (2005) Population Genetic 

structure of the Hawaiian precious 

coral Corallium lauuense (Octocorallia: 

Coralliidae) using microsatellites. Pages 

663–678 in Freiwald A, Roberts JM (eds.), 

Cold-water corals and ecosystems. 

Springer-Verlag Berlin Heidelberg

Barnes RD (1987) Invertebrate zoology, fifth 

edition. Saunders Publishing, New York.

Bates NR, Pequignet AC, Johnson RJ, Gruber N 

(2002) A short-term sink for atmospheric 

CO2 in subtropical mode water of the North 

Atlantic Ocean. Nature 420:489–493

Bayer FM (1956) Descriptions and re-descriptions 

of the Hawaiian Octocorals collected 

by the U.S. Fish Commission steamer 

Albatross 2. Gorgonacea: Scleraxonia. 

Pacific Science 10:67–95

Bayer FM (1981) Key to the genera of Octocorallia 

exclusive of Pennatulacea (Coelenterata: 

Anthozoa), with diagnosis of new taxa. 

Proceedings of the Biological Society of 

Washington 94:901–947

Bayer FM, Muzik KM (1977) An Atlantic helioporan 

coral (Coelenterata: Octocorallia). 

Proceeding of the Biological Society of 

Washington 90(4):975-984

Bayer EM, Stefani J (1987) New and previously 

known taxa of isidid octocorals 

(Coelenterata: Gorgonacea), partly from 

Antarctic waters, with descriptions of 

new taxa. Proceedings of the Biological 

Society of Washington 100(4):937–991

Bayer FM, Cairns SD (2003) A new genus 

of the scleraxonian family Coralliidae 

(Octocorallia: Gorgonacea) Proceedings 

of the Biological Society of Washington 

116(1):222–228

Bindoff NL, Willebrand J, Artale V, Cazenave A, 

Gregory J, Gulev S, Hanawa K, Le Quéré 

C, Levitus S, Nojiri Y, Shum CK, Talley 

LD, Unnikrishnan A (2007) Observations: 

oceanic climate change and sea level. 

Pages 385–432 in Solomon S, Qin D, 

Manning M, Chen Z, Marquis M, Averyt 

KB, Tignor M, Miller HL (eds.), Climate 

change 2007: The physical science 

basis. Contribution of Working Group I 

to the Fourth Assessment Report of the 

Intergovernmental Panel on Climate 

Change. Cambridge University Press, 

New York

Birkeland C (1974) Interactions between a sea 

pen and seven of its predators. Ecological 

Monographs 44:211–232

Boehlert GW, Genin A (1987) A review of the 

effects of seamounts on biological 

processes. Pages 319–334 in Keating 

BH, Fryer P, Batiza R, Boehlert GW 

(eds.), Seamounts, islands and atolls. 

Geophysical Monograph 43. American 

Geophysical Union, Washington, DC

Brancato MS, Bowlby CE (2005) Survey of 

fishing gear and fiber optic cable impacts 

to benthic habitats in the Olympic Coast 

National Marine Sanctuary. Pages 629–

630 in Barnes PW, Thomas JP (eds.), 

Benthic habitats and the effects of fishing. 

American Fisheries Society Symposium 

41, Bethesda, Maryland
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

50

Brancato MS, Bowlby CE, Hyland J, Intelmann 

SS, Brenkman K (2007) Observations of 

deep coral and sponge assemblages in 

Olympic Coast National Marine Sanctuary, 

Washington. Cruise Report: NOAA Ship 

McArthur II Cruise AR06-06/07. Marine 

Sanctuaries Conservation Series NMSP07-03. NOAA National Marine Sanctuary 

Program, Silver Spring, Maryland

Breeze H, Davis DS, Butler M, Vladmir K (1997) 

Distribution and status of deep sea corals 

off Nova Scotia. Marine Issues Committee 

Special Publication Number 1. Ecology 

Action Centre, Halifax, Nova Scotia

Broch H (1914) Stylasteridae. The Danish Ingolf 

Expedition 5(5):1–28

Brodeur RD (2001) Habitat-specific distribution of 

Pacific ocean perch (Sebastes alutus) in 

Pribilof Canyon, Bering Sea. Continental 

Shelf Research 21:207–224

Brooke S, Young CM (2003) Reproductive 

ecology of a deep-water scleractinian 

coral, Oculina varicosa, from the southeast 

Florida shelf. Continental Shelf Research 

23:847–858

Brooke S, Young CM (2005) Embryogenesis 

and larval biology of the ahermatypic 

scleractinian Oculina varicosa. Marine 

Biology 146(4):665–675

Buhl-Mortensen L, Mortensen PB (2005) 

Distribution and diversity of species 

associated with deep-sea gorgonian 

corals off Atlantic Canada. Pages 849–

879 in Freiwald A, Roberts JM (eds.), 

Cold-water corals and ecosystems. 

Springer-Verlag Berlin Heidelberg

Burgess SN, Babcock RC (2005) Reproductive 

ecology of three reef-forming, deep-sea 

corals in the New Zealand region. Pages 

701–713 in Freiwald A, Roberts JM (eds.), 

Cold-water corals and ecosystems. 

Springer-Verlag Berlin Heidelberg

Butler M, Gass S (2001) How to protect corals 

in Atlantic Canada. Pages 156–165 in 

Willison JHM et al. (eds.), Proceedings 

of the First International Symposium on 

Deep-Sea Corals. Ecology Action Centre 

and Nova Scotia Museum, Halifax, Nova 

Scotia.

Cairns SD (1979) The deep-water Scleractinia 

of the Caribbean Sea and adjacent 

waters. Pages 1–341 in Hummelinck PW, 

Van der Steen LJ (eds.), Studies on the 

fauna of Curacao and other Caribbean 

islands. Volume 57 Utrecht, Foundation 

for Scientific Research in Surinam and 

the Netherlands 

Cairns SD (1982) Antarctic and subantarctic 

Scleractinia. Antarctic Research Series 

34

Cairns SD (1984) New records of ahermatypic 

corals (Scleractinia) from the Hawaiian 

and Line islands. Occasional Papers 

of the Bernice Pauahi Bishop Museum 

25(10)

Cairns SD (1992a) A generic revision of the 

Stylasteridae. Part 3. Keys to the genera. 

Bulletin of Marine Science 49:538–545

Cairns SD (1992b) Worldwide distribution of the 

Stylasteridae. Scientia Marina 56:125–

130

Cairns SD (1995) The marine fauna of New 

Zealand: Scleractinia (Cnidaria: Anthozoa) 

New Zealand Oceanographic Institute 

Memoir 103:1–210

Cairns SD (2001) A brief history of taxonomic 

research on azooxanthellate Scleractinia 

(Cnidaria: Anthozoa) Bulletin of the 

Biological Society of Washington 10:191–

203

Cairns SD (2006) New records of azooxanthellate 

Scleractinia from the Hawaiian Islands. 

Occasional Papers of the Bernice Pauahi 

Bishop Museum 87:45–53

Cairns SD (In press) Deep-water corals: An 

overview with special reference to 

diversity and distribution of deep-water 

scleractinian corals. Bulletin of Marine 

Science 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

51 INTRODUCTION AND OVERVIEW

Cairns S, Stanley G (1982) Ahermatypic coral 

banks: Living and fossil counterparts. 

Proceedings of the Fourth International 

Coral Reef Symposium, Manila. 1:611–

618

Cairns, SD, Hoeksema BW, van der Land J (1999) 

List of extant stony corals. Appendix pp. 

13-46 In Cairns SD, Species richness 

of Recent Scleractinia. Atoll Research 

Bulletin 459:1-46

Cairns SD, Bayer FM (2002) Studies on western 

Atlantic Octocorallia (Coelenterata: 

Anthozoa) Part 2: The genus Callogorgia

Gray, 1858. Proceedings of the Biological 

Society of Washington 115(4):840–867

Cairns SD, Bayer FM (2005) A review of the genus 

Primnoa (Octocorallia: Gorgonacea: 

Primnoidae), with the description of two 

new species. Bulletin of Marine Science 

77(2):225–256

Caldeira K, Wickett ME (2003) Oceanography: 

Anthropogenic carbon and ocean pH. 

Nature 425:365

Caldeira K, Wickett ME (2005) Ocean model 

predictions of chemistry changes 

from carbon dioxide emissions to the 

atmosphere and ocean. Journal of 

Geophysical Research–Oceans 110(c9):

C09S04

Chuenpagdee, R, Morgan LE, Maxwell SM, 

Norse EA, Pauly D (2003) Shifting gears: 

assessing collateral impacts of fishing 

methods in U.S. waters. Frontiers in 

Ecology and the Environment 1(10):517–

524

Costello MJ, McCrea M, Freiwald A, Lundalv T, 

Jonsson L, Bett BJ, Weering TV, de Haas 

H, Roberts JM, Allen D (2005) Functional 

role of deep-sea cold-water Lophelia coral 

reefs as fish habitat in the north-eastern 

Atlantic. Pages 771–805 in Freiwald A, 

Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Dawson J (2002) Biogeography of azooxanthellate 

corals in the Caribbean and surrounding 

areas. Coral Reefs 21(1):27–40

Denman KL, Brasseur G, Chidthaisong A, Ciais P, 

Cox PM, Dickinson RE, Hauglustaine D, 

Heinze C, Holland E, Jacob D, Lohmann 

U, Ramachandran S, da Silva Dias PL, 

Wofsy SC, Zhang X (2007) Couplings 

between changes in the climate system 

and biogeochemistry. Pages 499–588 in 

Solomon S, Qin D, Manning M, Chen Z, 

Marquis M, Averyt KB, Tignor M, Miller HL 

(eds.), Climate change 2007: the physical 

science basis. Contribution of Working 

Group I to the Fourth Assessment Report 

of the Intergovernmental Panel on Climate 

Change. Cambridge University Press, 

New York

DeVogeleare A, Burton EJ, Trejo T, King CE, 

Clague DA, Tamburri MN, Cailliet GM, 

Kochevar RE, Douros WJ (2005) Deepsea corals and resource protection at 

the Davidson Seamount, California, 

USA Pages 1189–1198 in Freiwald A, 

Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Druffel ERM, Griffin S, Witter A, Nelson E, 

Southon J, Kashgarian M, Vogel J (1995) 

Gerardia: bristlecone pine of the deepsea? Geochimica et Cosmochimica Acta 

59(23):5031–5036

Ehrlich, H., Etnoyer P, Litvinov SD, Olennikova M, 

Domaschke H, Hanke T, Born R, Meissner 

H, Worch H (2006) Biomaterial structure 

in deep-sea bamboo coral (Gorgonacea: 

Isididae): perspectives for the development 

of bone implants. Materialwissenschaft 

und Werkstofftechnik 37(6):553–557

Emeis KC, Benoit JR, Deegan L, Gilbert AJ, 

Lee V, Glade JM, Meybeck M, Olsen 

SB, von Bodungen B (2001) Group 4: 

unifying concepts for integrated coastal 

management. Page 378 in von Bodungen 

G, Turner K (eds.), Science and integrated 

coastal management. Dahlem University 

Press, Berlin

Etnoyer P, Morgan L (2003) Occurrences of 

habitat-forming deep sea corals in the 

northeast Pacific Ocean. A report to 

NOAA’s Office of Habitat Conservation. 

Marine Conservation Biology Institute, 

Redmond, Washington
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

52

Etnoyer P, Morgan L (2005) Habitat-forming 

deep-sea corals in the Northeast Pacific 

Ocean. Pages 331–343 in Freiwald A, 

Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Etnoyer P, Warenchuk J (in press) On the 

occurrence of a shark nursery (family 

Seyliorhinidae) in the deep-water 

gorgonian field in the Mississippi Canyon, 

Gulf of Mexico. Bulletin of Marine Science 

80(2)

Etnoyer P, Cairns SD, Sanchez JA, Reed JK, 

Lopez JV, Schroeder WW, Brooke SD, 

Watling L, Baco-Taylor A, Williams 

GC, Lindner A, France SC, Bruckner 

AW (2006) Deep-sea coral collection 

protocols. NOAA Technical Memorandum 

NMFS-OPR-28. Silver Spring, Maryland

Fabricius K, Alderslade P (2001) Soft corals and 

sea fans. Australian Institute of Marine 

Science, Townsville, Australia

Feely R, Sabine C (2004) Impact of anthropogenic 

CO2 on the CaCO3 system in the oceans. 

Science 305:362–366

Food and Agriculture Organization of the United 

Nations (FAO) (2005) Impact of trawling 

and scallop dredging on benthic habitats 

and communities. FAO Fisheries 

Technical Paper 472. Rome

Fosså, JH, Mortensen PB, Furevik DM (2002) 

The deep-water coral Lophelia pertusa in 

Norwegian waters: distribution and fishery 

impacts. Hydrobiologia 471:1–12.

Frederiksen R, Jensen A, Westerberg H (1992) 

The distribution of the scleractinian 

coral Lophelia pertusa around the Faroe 

Islands and the relation to internal tidal 

mixing. Sarsia 77:157–171. 

Freese JL, Auster PJ, Heifetz J, Wing BL (1999) 

Effects of trawling on seafloor habitat and 

associated invertebrate taxa in the Gulf of 

Alaska. Marine Ecology Progress Series 

182:119–126.

Freiwald A, Fosså JH, Grehan A, Koslow T, 

Roberts JM (2004) Cold-water coral reefs. 

United Nations Environment Programme 

- World Conservation Monitoring Centre. 

Cambridge, UK

French LS, Kazanis EG, Labiche LC, Montgomery 

TM, Richardson GE (2005) Deepwater 

Gulf of Mexico 2005: interim report of 2004 

highlights. MMS report OCS 2005-023. 

U.S. Department of the Interior, Minerals 

Management Service, New Orleans.

Gass SE, Roberts JM (2006) The occurrence 

of the cold-water coral Lophelia pertusa 

(Scleractinia) on oil and gas platforms in 

the North Sea: colony growth, recruitment 

and environmental controls on distribution. 

Marine Pollution Bulletin 52:549–559.

Gass SE, Willison JHM (2005) An assessment 

of the distribution of deep-sea corals in 

Atlantic Canada by using both scientific 

and local forms of knowledge. Pages 

223–245 in Freiwald A, Roberts JM (eds.), 

Cold-water corals and ecosystems. 

Springer-Verlag Berlin Heidelberg

Gattuso JP, Allemand D, Frankignoulle M (1999) 

Photosynthesis and calcification at 

cellular, organismal and community level 

in coral reefs: a review on interactions and 

control by carbonate chemistry. American 

Zoologist 39:160–183.

George RY (2004a) Conservation and 

management of deep-sea coral reefs: 

need for new definition, an agenda and 

action plan for protection. Deep-Sea 

Newsletter 33:25–29.

George RY (2004b) Lophelia bioherms and 

lithoherms as fish habitats on the Blake 

Plateau: biodiversity and sustainability. 

ICES Annual Science Conference 

Proceedings, Vigo, Spain, September 

24–26, 2004. <www.ices.dk/iceswork/

asc/2004/abstracts/abstracts/AA.pdf>

Grange KR. (1985) Distribution, standing crop, 

population structure and growth rates of 

an unexploited resource of black coral 

in the southern fjords of New Zealand. 

Proceedings of the Fifth International 

Coral Reef Congress, Tahiti, May 27–

June 1, 1985. 6:217–222. 
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

53 INTRODUCTION AND OVERVIEW

Grange KR (1997) Demography of black coral 

populations in Doubtful Sound, New 

Zealand: Results from a 7-year experiment. 

Pages 185–193 in Proceedings of the 6th 

International Conference on Coelenterate 

Biology 1995. National Museum of Natural 

History, Leiden

Grasshoff M (1979) Zur bipolaren verbreitung der Zur bipolaren verbreitung der

oktocoralle Paragorgia arborea (Cnidaria: 

Anthozoa:Scleraxonia) Senckengergiana 

Maritima 11:115–137.

Grehan A, Unnithan V, Wheeler A, Monteys X, 

Beck T, Wilson M, Guinan J, Foubert A, 

Klages M, Thiede J (2004) Evidence 

of major fisheries impact on cold-water 

corals in the deep waters off the Porcupine 

Bank, West Coast of Ireland: Are interim 

management measures required? <www.

ices.dk/iceswork/asc/2004/abstracts/

abstracts/AA.pdf>

Grehan AJ, Unnithan V, Olu-Le Roy K, 

Opderbecke J (2005) Fishing impacts 

on Irish deepwater coral reefs: Making a 

case for coral conservation. Pages 819–

832 in Barnes PW, Thomas JP (eds.), 

Benthic habitats and the effects of fishing. 

American Fisheries Society Symposium 

41, Bethesda, Maryland

Grigg RW (1974) Distribution and abundance 

of precious corals in Hawaii. Pages 235-

240 in Cameron AM (ed.) Proceedings 

of the Second International Coral Reef 

Symposium Brisbane Reef, Australia Vol. 

II:

Grigg RW (1976) Fishery management of 

precious and stony corals in Hawaii. Sea 

Grant Technical Report UNIHI-SeagrantTR-77-03.48 University of Hawaii 48 pp.

Grigg RW (1993) Precious coral fisheries of 

Hawaii and the U.S. Pacific Islands. 

Marine Fisheries Review 55:50–60

Grigg R (2002) Precious corals in Hawaii: 

discovery of a new bed and revised 

management measures for existing beds. 

Marine Fisheries Review 64:13–20

Grigg RW (2003) Invasion of a deep black coral 

bed by an alien species, Carijoa riisei, off 

Maui, Hawaii. Coral Reefs 22:121–122

Guinotte JM, Orr J, Cairns S, Freiwald A, Morgan 

L, George R (2006) Will human-induced 

changes in seawater chemistry alter the 

distribution of deep-sea scleractinian 

corals? Frontiers in Ecology 4(3):141–

146

Hall-Spencer J, Allain V, Fosså JH (2001) Trawling 

damage to northeast Atlantic ancient coral 

reefs. Proceedings of the Royal Society 

of London Series B 269:507–511

Heifetz J (2002) Coral in Alaska: distribution, 

abundance and species associations. 

Hydrobiologia 471:19–28

Heifetz J, Wing BL, Stone RP, Malecha PW, 

Courtney DL (2005) Corals of the 

Aleutian Islands. Fisheries Oceanography 

14(Suppl. 1):131–138

High WL (1998) Observations of a scientist/

diver on fishing technology and fisheries 

biology. Alaska Fisheries Science Center 

Processed Report 98-01. NOAA National 

Marine Fisheries Service, Seattle

Husebø A, Nøttestad L, Fosså JH, Furevik DM, 

Jørgensen SB (2002) Distribution and 

abundance of fish in deep-sea coral 

habitats. Hydrobiologia 471:91–99

Hutchings JA (2000) Collapse and recovery of 

marine fishes. Nature 406:882–885

Hyland J, Cooksey C, Bowlby E, Brancato MS, 

Intelmann S (2005) A pilot survey of 

deepwater coral/sponge assemblages 

and their susceptibility to fishing/harvest 

impacts at the Olympic Coast National 

Marine Sanctuary (OCNMS) Cruise 

Report for NOAA Ship McARTHUR II 

Cruise AR-04-04: Leg 2. NOAA Technical 

Memorandum NOS NCCOS 15. NOAA/

NOS Center for Coastal Environmental 

Health and Biomolecular Research, 

Charleston, South Carolina

International Council on the Exploration of the 

Seas (ICES) (2005) Report of the Working 

Group on Deep-water Ecology. ICES CM 

2005/ACE:02, Copenhagen
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

54

Intergovernmental Panel on Climate Change 

(IPCC) (2001) Climate change 2001: the 

scientific basis. Cambridge University 

Press, New York

IPCC (2005) Carbon dioxide capture and storage: 

summary for policymakers. Special 

Report. Eighth session of IPCC Working 

Group III, September 22–24, 2005, 

Montreal, Canada

IPCC (2007a) Summary for policymakers in 

Solomon S, Qin D, Manning M, Chen Z, 

Marquis M, Averyt KB, Tignor M, Miller HL 

(eds.), Climate change 2007: the physical 

science basis. Contribution of Working 

Group I to the Fourth Assessment Report 

of the Intergovernmental Panel on Climate 

Change. Cambridge University Press, 

New York

IPCC (2007b) Summary for policymakers. Pages 

7–22 in Parry ML, Canziani OF, Palutikof 

JP, van der Linden PJ, Hanson CE (eds.), 

Climate change 2007: impacts, adaptation 

and vulnerability. Contribution of Working 

Group II to the Fourth Assessment Report 

of the Intergovernmental Panel on Climate 

Change. Cambridge University Press, 

New York

International Seabed Authority (ISA) (2007) 

Polymetallic sulfides and cobalt crusts: 

Their environment and considerations 

for establishment of environmental 

baselines associated with monitoring 

programmes for exploration. Available: Available:

http://www.isa.org.jm/files/documents/

EN/Workshops/2004/Proceedings-ae.pdf

Joos F, Plattner GK, Stocker TF, Marchal O, 

Schmittner A (1999) Global warming and 

marine carbon cycle feedbacks on future 

atmospheric CO2. Science 128:464–467

Kastendiek J (1976) Behavior of the sea pansy 

Renilla kollikeri Pfeffer (Coelenterata: 

Pennatulacea) and its influence on the 

distribution and biological interactions of 

the species. Biological Bulletin 151:518–

537

Kelley C, Ikehara W (2006) The impacts of 

bottomfishing on Raita and West St. 

Rogatien Banks in the Northwestern 

Hawaiian Islands. Atoll Research Bulletin 

543:305–317

Kleypas JA, Buddemeier RW, Archer D, Gattuso 

J, Landon C, Opdyke BN (1999) 

Geochemical consequences of increased 

carbon dioxide in coral reefs. Science 

284:118–120

Kleypas JA, Feely RA, Fabry VJ, Langdon C, 

Sabine CL, Robbins LL (2006) Impacts 

of ocean acidification on coral reefs and 

other marine calcifiers: a guide for future 

research. Report of a workshop held April 

18–20, 2005, St. Petersburg, Florida

Koenig CC (2001) Oculina Banks: habitat, fish 

populations, restoration and enforcement. 

Report to the South Atlantic Fishery 

Management Council. Http://www.safmc.

net 

Koenig, CC, Coleman F, Scanlon KM (2000) 

Reef fish resources and gas pipelines 

on the West Florida Shelf: potential 

conflict. Pages 109–119 in Vigil D (ed.), 

Proceedings: Gulf of Mexico fish and 

fisheries: bringing together new and 

recent research. MMS report October 

2000

Koenig CC, Shephard AN, Reed JK, Coleman 

FC, Brooke SD, Brusher J, Scanlon KM 

(2005) Habitat and fish populations in 

the deep-sea Oculina coral ecosystem of 

the Western Atlantic. Pages 795–805 in 

Barnes PW, Thomas JP (eds.), Benthic 

habitats and the effects of fishing. 

American Fisheries Society Symposium 

41, Bethesda, Maryland

Koslow JA, Boehlert GW, Gordon JDM, Haedrich 

RL, Lorance P, Parin N (2000) Continental 

slope and deep-sea fisheries: implications 

for a fragile ecosystem. ICES Journal of 

Marine Science 57:548–557

Koslow J, Gowlett-Holmes K, Lowry J, Poore 

G, Williams A (2001) Seamount benthic 

macrofauna off southern Tasmania: 

Community structure and impacts of 

trawling. Marine Ecology Progress Series 

213:111–125
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

55 INTRODUCTION AND OVERVIEW

Krieger K (1998) Primnoa spp. observed inside 

and outside a bottom trawl path from a 

submersible. Abstract. 10th Western 

Groundfish Conference

Krieger, KJ (2001) Coral (Primnoa) impacted 

by fishing gear in the Gulf of Alaska. 

Pages 106–116 in Willison et al. (eds.), 

Proceedings of the First International 

Symposium on Deep-Sea Corals, Ecology 

Action Centre and Nova Scotia Museum, 

Halifax, Nova Scotia

Krieger KJ, Wing BL (2002) Megafaunal 

associations with deepwater corals 

(Primnoa spp.) in the Gulf of Alaska. 

Hydrobiologia 471:83–90

Langton RW, Langton EW, Theroux RB, Uzmann 

JR (1990) Distribution, behavior and 

abundance of sea pens, Pennatula 

aculeate, in the Gulf of Maine. Marine 

Biology 107:463–469

Le Goff-Vitry M, Pybus OG, Rogers AD (2004) 

Genetic structure of the deep-sea coral 

Lophelia pertusa in the northeast Atlantic 

revealed by microsatellites and internal 

transcribed spacer sequences. Molecular 

Ecology 13:537–549

Le Goff-Vitry MC, Rogers AD (2005) Molecular 

ecology of Lophelia pertusa in the NE 

Atlantic. Pages 653–662 in Freiwald A, 

Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Leverette TL, Metaxas A (2005) Predicting 

habitat for two species of deep water coral 

on the Canadian Atlantic continental shelf 

and slope. Pages 467–479 in Freiwald 

A, Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Love MS, Yoklavich MM, Black BA, Andrews AH 

(2007) Age of black coral (Antipathes 

dendrochristos Opresko, 2005) colonies, 

with notes on associated invertebrate 

species. Bulletin of Marine Science 

80(2):391–400

Maragos JE, Potts DC, Aeby G, Gulko D, Kenyon 

J, Siciliano D, VanRavenswaay D (2004) 

2000–2002 rapid ecological assessment 

of corals (Anthozoa) on shallow reefs 

of the Northwestern Hawaiian Islands. 

Part 1: species and distribution. Pacific 

Science 58:211–230

McDonough JJ, Puglise KA (2003) Summary: 

Deep-Sea Corals Workshop. International 

planning and collaboration workshop for 

the Gulf of Mexico and the North Atlantic 

Ocean. Galway, Ireland, January 16-

17, 2003. U.S. Department. Commerce, 

NOAA Technical. Memorandum NMFS-F/

SPO-60, 51 pp 

Messing, CG, Neuman AC, Lang JC (1990) 

Biozonation of deep-water lithoherms 

and associated hardgrounds in the 

northeastern Straits of Florida. Palaios 

5:15–33.

Miller, KJ. 1996. Piecing together the reproductive 

habitats of New Zealands endemic black 

corals. Water Atmosphere. 4:18-19 

Miller JE, Hoeke RK, Appelgate TB, Johnson PJ, 

Smith J, Bevacqua S (2003) Bathymetric 

atlas of the Northwestern Hawaiian 

Islands, draft. National Oceanic and 

Atmospheric Administration and Hawaii 

Mapping Research Group.

Minerals Management Service (2003) Grid 

ecological assessment and ROV survey 

status report. Available: www.gomr.mms. Available: www.gomr.mms.

gov/homepg/regulate/environ/ea_grid/

ea_grid.asp.

Morgan LE, Tsao C, Guinotte JM (2006) Status 

of deep sea corals in U.S. waters with 

recommendations for their conservation 

and management. Marine Conservation 

Biology Institute, Bellevue, Washington

Mortensen PB (2000) Lophelia pertusa in 

Norwegian waters: distribution, growth and 

associated fauna. Doctoral dissertation, 

University of Bergen, Department of 

Fisheries and Marine Biology
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

56

Mortensen PB (2001) Aquarium observations on 

the deep-water coral Lophelia pertusa 

(L., 1758) (Scleractinia) and selected 

associated invertebrates. Ophelia 

54(2):83–104

Mortensen PB, Buhl-Mortensen L (2005a) Coral 

habitats in The Gully, a submarine canyon 

off Atlantic Canada. Pages 247–277 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin Heidelberg

Mortensen PB, Buhl-Mortensen L (2005b) 

Morphology and growth of the deepwater 

gorgonians Primnoa resedaeformis and 

Paragorgia arborea. Marine Biology 

147:775–788

Mortensen PB, Fosså JH (2006) Species diversity 

and spatial distribution of invertebrates 

on Lophelia reefs in Norway. Pages 

1849–1868 in Proceedings of the 10th

International Coral Reef Symposium. 

Okinawa, Japan

Mortensen PB, Rapp HT (1998) Oxygen and 

carbon isotope ratios related to growth 

line patterns in skeletons of Lophelia 

pertusa (L) (Anthozoa, Scleractinia): 

implications for determination of linear 

extension rates. Sarsia 83:433–446

Mortensen PB, Buhl-Mortensen L, Gordon Jr. 

DC, Fader GBJ, McKeown DL, Fenton 

DG (2005) Effects of fisheries on deepwater gorgonian corals in the Northeast 

Channel, Nova Scotia (Canada) American 

Fisheries Society Symposium 41:369–

382

National Research Council (2002) Effects of 

trawling and dredging on seafloor habitat. 

National Academy Press, Washington, 

DC

Newton CR, Mullins HT, Gardulski AF (1987) 

Coral mounds on the West Florida Slope: 

unanswered questions regarding the 

development of deepwater banks. Palaios 

2:359–367

Opresko, DM (2001) Revision of the Antipatharia 

(Cnidaria: Anthozoa) Part I. Establishment 

of a new family, Myriopathidae. Zoologische 

Mededelingen, Leiden 75:147–174

Opresko, DM (2002) Revision of the 

Antipatharia (Cnidaria: Anthozoa) 

Part II. Schizopathidae. Zoologische 

Mededelingen, Leiden 76:411–442

Opresko, DM (2003) Revision of the 

Antipatharia (Cnidaria: Anthozoa) 

Part III. Cladopathidae. Zoologische 

Mededelingen, Leiden 77:495–536

Opresko, DM (2004) Revision of the Antipatharia 

(Cnidaria: Anthozoa, Antipatharia) 

Part IV. Establishment of a new 

family, Aphanipathidae. Zoologische 

Mededelingen, Leiden 78:209–240

Ostarello GL (1973) Natural history of the 

hydrocoral Allopora californica (Verrill 

1866) Biological Bulletin 145:548–564

Ostarello GL (1976) Larval dispersal in the 

subtidal hydrocoral Allopora californica 

(Verrill 1866) Pages 331–337 in Mackie 

GO (ed.), Coelenterate ecology and 

behavior. Plenum Press, New York

Parker NR, Mladenov PV, Grange KR (1997) 

Reproductive biology of the antipatharian 

black coral Antipathes fordensis in 

Doubtful Sound, Fjordland, New Zealand. 

Marine Biology 130:11–22

Pourtalès LF de (1867) Contributions to the 

fauna of the Gulf Stream at great depths. 

Bulletin of the Museum of Comparative 

Zoology 1(6):103–120

Pourtalès LF de (1871) Deep-sea corals. 

Illustrated catalogue of the Museum of 

Comparative Zoology at Harvard College 

4 Probert PK, McKnight DG, Grove SL 

(1997) Benthic invertebrate bycatch from 

a deepwater trawl fishery, Chatham Rise, 

New Zealand. Aquatic Conservation: 

Marine and Freshwater Ecosystems 

7:27–40
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

57 INTRODUCTION AND OVERVIEW

Puglise KA, Brock RJ, McDonough III JJ (2005) 

Identifying critical information needs and 

developing institutional partnerships to 

further the understanding of Atlantic deepsea coral ecosystems. Pages 1129–1140 

in Freiwald A, Roberts JM (eds.), Coldwater corals and ecosystems. SpringerVerlag Berlin Heidelberg

Reed JK (1980) Distribution and structure of 

deep-water Oculina varicosa coral reefs 

off central eastern Florida. Bulletin of 

Marine Science 30(3):667–677

Reed JK (1981) In situ growth rates of the 

scleractinian coral Oculina varicosa

occurring with zooxanthellae on 6 m reefs 

and without on 80 m banks. Proceedings 

of the Fourth International Coral 

Reef Symposium, May 1981, Manila, 

Philippines 2:201–206

Reed JK (2002a) Comparison of deep-water 

coral reefs and lithoherms off southeastern 

U.S.A. Hydrobiologia 471:57–69

Reed JK (2002b) Deep-water Oculina coral 

reefs of Florida: biology, impacts, and 

management. Hydrobiologia 471:43–55

Reed JK, Shepard A, Koenig C, Scanlon K, 

Gilmore G (2005) Mapping, habitat 

characterization, and fish surveys of the 

deep-water Oculina coral reef Marine 

Protected Area: a review of historical 

and current research. Pages 443–465 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin Heidelberg

Reed JK, Weaver D, Pomponi SA (2006) Habitat 

and fauna of deep-water Lophelia pertusa

coral reefs off the Southeastern USA: 

Blake Plateau, Straits of Florida, and Gulf 

of Mexico. Bulletin of Marine Science 

78(2):343–375

Reyes J, Santodomingo N, Gracia A, BorreroPérez G, Navas G, Ladino LM, Bermúdez A, 

Benavides M (2005) Southern Caribbean 

azooxanthellate coral communities off 

Colombia. Pages 309-330 in Freiwald 

A, Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Richer de Forges B, Koslow JA, Poore GCB 

(2000) Diversity and endemism of the 

benthic seamount fauna in the southwest 

Pacific. Nature 405:944–947

Risk MJ, MacAllister DE, Behnken L (1998) 

Conservation of cold water warm water 

seafans: threatened ancient gorgonian 

groves. Sea Wind 12(1):2–21

Risk MJ, Heikoop JM, Snow MG, Beukens B 

(2002) Lifespans and growth patterns 

of two deep sea corals: Primnoa 

resedaeformis and Desmophyllum 

cristagalli. Hydrobiologia 471:125–131

Roark EB, Guilderson TP, Dunbar RB, Ingram 

BL (2006) Radiocarbon-based ages 

and growth rates of Hawaiian deep-sea 

corals. Marine Ecology Progress Series 

327:1–14

Roberts CM (2002) Deep impact: the rising toll of 

fishing in the deep sea. Trends in Ecology 

and Evolution 17:242–245

Roberts JM, Long D, Wilson JB, Mortensen 

PB, Gage JD (2003) The coldwater 

coral Lophelia pertusa (Scleractinia) 

and enigmatic seabed mounds along 

the north-east Atlantic margin: are they 

related? Marine Pollution Bulletin 46:7–

20

Roberts JM, Brown CJ, Long D, Bates CR (2005) 

Acoustic mapping using a multibeam 

echosounder reveals cold-water coral 

reefs and surrounding habitats. Coral 

Reefs 24:654–669

Roberts JM, Wheeler AJ, Freiwald A (2006) Reefs 

of the deep: The biology and geology of 

cold-water coral ecosystems. Science 

312:543–547

Rogers AD (1999) The biology of Lophelia pertusa

(Linnaeus 1758) and other deep-water 

reef-forming corals and impacts from 

human activities. International Review of 

Hydrobiology 84:315–406
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

58

Rogers AD (2004) The biology, ecology and 

vulnerability of seamount communities. 

International Union for Conservation 

of Nature and Natural Resources, <

http://www.iucn.org/themes/marine/pdf/

AlexRogers-CBDCOP7-SeamountsComplete1.pdf>

Royal Society (2005) Ocean acidification due to 

increasing atmospheric carbon dioxide. 

The Royal Society Policy document 12/05, 

June 2005. Clyvedon Press, Cardiff, UK

Ryland JS, Babcock RC (1991) Annual cycle of 

gametogenesis and spawning in a tropical 

zoanthid, Protopalythoa sp. Hydrobiologia 

(216/217):117-123 

Sabine CL, Feely RA, Gruber N, Key RM, Lee 

K, Bullister JL, Wanninkhof R, Wong CS, 

Wallace DWR, Tilbrook B, Millero FJ, 

Peng T-H, Kozyr A, Ono T, Rios AF (2004) 

The Oceanic Sink for Anthropogenic CO2. 

Science 305(5682), 367-371

Santangelo G, Carlietti E, Maggi E, Bramanti 

L (2003) Reproduction and population Reproduction and population 

sexual structure of the overexploited 

Mediterranean red coral Corallium rubrum. 

Marine Ecology Progress Series 248:99–

108

Schroder-Ritzrau A, Freiwald A, Mangini A (2005) 

U/Th dating of deep-water corals from the 

eastern North Atlantic and the western 

Mediterranean Sea. Pages 157–172 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin Heidelberg

Schroeder WW (2002) Observations of Lophelia 

pertusa and the surficial geology at a 

deepwater site in the northeastern Gulf of 

Mexico. Hydrobiologia 471:29–33

Schrope, M (2007) Digging deep. Nature 

447:246–247

Smith N (1983) Temporal and spatial 

characteristics of summer upwelling 

along Florida’s Atlantic shelf. Journal of 

Physical Oceanography 13:1709–1715

Smith PJ (2001) Managing biodiversity: 

invertebrate by-catch in seamount 

fisheries in the New Zealand exclusive 

economic zone (a case study) United 

Nations Environment Programme: 

Workshop on Managing Global Fisheries 

for Biodiversity, Victoria

Smith RC, Ainley D, Baker K, Domack E, Emslie S, 

Fraser B, Kennett J, Leventer A, MosleyThompson E, Stammerjohn S, Vernet M 

(1999) Marine ecosystem sensitivity to 

climate change. Bioscience 49:393–404

Stetson TR, Squires DF, Pratt RM (1962) Coral 

banks occurring in deep water on the 

Blake Plateau. American Museum of 

Natural History 2113:1–39

Stone RP (2006) Coral habitat in the Aleutian 

Islands of Alaska: depth distribution, finescale species association, and fisheries 

interactions. Coral Reefs 25:229–238

Strychar KB, Hamilton LC, Kenchington EL, Scott 

DB (2005) Genetic circumscription of 

deep-water coral species in Canada using 

18S rRNA. Pages 679–690 in Freiwald 

A, Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Theroux RB, Wigley RL (1998) Quantitative 

composition and distribution of the 

macrobenthic invertebrate fauna of 

the continental shelf ecosystems of 

the northeastern United States. NOAA 

Technical Report NMFS-140. Silver 

Spring, Maryland

Tissot BN, Yoklavich MM, Love MS, York K, Amend 

M (2006) Benthic invertebrates that 

form habitat on deep banks off southern 

California, with special reference to deep 

sea coral. Fishery Bulletin 104(2):167–

181

Torrents O, Garrabou J, Marschal C, Hamelin JG 

(2005) Age and size at first reproduction 

in the commercially exploited red coral 

Corallium rubrum (L.) in the Marseilles 

area (France, NW Mediterranean) 

Biological Conservation 121:391–397
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

59 INTRODUCTION AND OVERVIEW

Uchupi E (1968) Atlantic continental shelf and 

slope of the United States—physiography. 

Geological Survey Professional Paper 

529-C:1–30

U.S. Coral Reef Task Force (2000) National 

action plan to conserve coral reefs. 

Washington, DC

Vaughan TW (1907) Recent Madreporaria of the 

Hawaiian Islands and Laysan. Bulletin 

United States National Museum 59

Vaughan TW, Wells JW (1943) Revision of 

the suborders, families and genera of 

the Scleractinia. Geological Society of 

America, Special. Papers, 44:1–363

Verrill AE (1883) Report on the Anthozoa, and 

on some additional material dredged by 

the Blake in 1877–1879, and by the U.S. 

Fish Commission Steamer “Fish Hawk.” 

Pages 1–72 in 1880-82: Bulletin of the 

Museum of Comparative Zoology at 

Harvard College 11

Verrill AE (1902) Papers on corals. Transactions 

of the Connecticut Academy of Arts and 

Sciences 11:63–266

Vitousek PM, D’Antonio CM, Loope LL, 

Westbrooks R (1996) Biological 

invasions as global environmental 

change. American Scientist 84:468–478

Waddell JE (ed.) (2005) The state of coral reef 

ecosystems of the United States and 

Pacific Freely Associated States: 2005. 

NOAA Technical Memorandum NOS 

NCCOS 11. NOAA/NCCOS Center for 

Coastal Monitoring and Assessment’s 

Biogeography Team. Silver Spring, 

Maryland

Waller RG, Tyler PA (2005) The reproductive 

biology of two deep-water, reef-building 

scleractinians from the NE Atlantic Ocean. 

Coral Reefs 24:514–522

Watling L, Auster P (2005) Distribution of deepwater Alcyonacea off the northeast coast 

of the United States. Pages 279–296 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin Heidelberg

Watling L, Auster P, Babb I, Skinder C, Hecker 

B (2003) A geographic database 

of deepwater alcyonaceans of the 

northeastern U.S. continental shelf and 

slope. Version 1.0 CD-ROM. National 

Undersea Research Center, University of 

Connecticut, Groton

Weinberg S (1976) Revision of the common 

Octocorallia of the Mediterranean 

circalittoral. I. Gorgonacea. Beaufortia 

24:63–104

Wheeler AJ, Bett BJ, Billett DS, Masson DG, 

Mayor D (2005) The impact of demersal 

trawling on northeast Atlantic coral 

habitats: The case of the Darwin Mounds, 

United Kingdom. Pages 807–817 in 

Barnes PW, Thomas JP (eds.), Benthic 

habitats and the effects of fishing. 

American Fisheries Society Symposium 

41, Bethesda, Maryland

Wilkinson C (2004) Status of coral reefs of the 

world: 2004. Global Coral Reef Monitoring 

Network, International Coral Reef 

Initiative, Australian Institute of Marine 

Science, Australia

Williams B, Risk MJ, Ross SW, Sulak KJ (2006) 

Deep-water Antipatharians: proxies 

of environmental change. Geology 

34(9):773–776

Williams B, Risk MJ, Ross SW, Sulak KJ (in press) 

Stable isotope records from deepwater 

Antipatharians: 400-year records from the 

south-eastern coast of the United States 

of America. Bulletin of Marine Science

Williams GC (1995) Living genera of sea 

pens (Coelenterata: Octocorallia: 

Pennatulacea): illustrated key and 

synopses. Zoological Journal of the 

Linnean Society 113:93–140

Williams GC (1999) Index Pennatulacea: 

annotated bibliography and indexes of 

the sea pens (Coelenterata: Octocorallia) 

of the world 1469–1999. Proceedings 

of the California Academy of Sciences

51(2):19–103
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

60

Wilson MT, Andrews AH, Brown AL, Cordes EE 

(2002) Axial rod growth and age estimation 

of the sea pen, Halipteris willemoesi

Kölliker. Hydrobiology 47:133–142 Hydrobiology 47:133–142

Zibrowius H (1980) Les Scléractiniaires de 

la Méditerranée et de l’Atlantique 

nord-oriental. Mémoires de l’Institut Mémoires de l’Institut 

Océanographique, Monaco, 11
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

61 INTRODUCTION AND OVERVIEW

Higher Taxon Species Alaska West Coast Pacific Islands

Phylum Cnidaria

Class Anthozoa

Subclass Hexacorallia

Order Scleractinia

Family 

Caryophylliidae Lophelia pertusa ●

Family 

Dendrophylliidae Enallopsammia rostrata ●

Dendrophyllia oldroydae ●

Family Oculinidae Oculina profunda ●

Order Antipatharia

Family Antipathidae Antipathes dendrochristos ●

Antipathes spp.* ●

Family Cladopathidae Chrysopathes formosa ● ○

Chrysopathes speciosa ● ○

Family 

Schizopathidae Bathypathes patula ● ○

Bathypathes sp. ●

Dendrobathypathes 

boutillieri ●

Order Zoanthidea

Family Gerardiidae Gerardia sp. ●

Subclass Octocorallia 

Order Alcyonacea

Family Neptheidae Eunephthea rubiformis ● ○

Order Gorgonacea

Family Coralliidae Corallium secundum ●

Corallium laauense ●

Family Isididae Isidella spp.* ● ● ●

Keratoisis profunda ●

Keratoisis sp.* ○ ● ○

Lepidisis sp.* ● ○ ○

Family Paragorgiidae Paragorgia arborea ● ●

Paragorgia sp.* ○

Family Primnoidae Fanellia sp.* ● ○ 

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, ○ 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
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

62

Higher Taxon Species Alaska West Coast Hawaii

Plumarella sp.* ● ○ ○

Primnoa pacifica ● ●

Order Pennatulacea

Family Anthoptilidae Anthoptilum spp. ● ○ ○

Family Halipteridae Halipteris willimoesi ● ○

Family Protoptilidae Protoptilum sp. ●

Class Hydrozoa 

Order Anthoathecatae

Family Stylasteridae Stylaster cancellatus ●

Stylaster campylecus ●
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

63 INTRODUCTION AND OVERVIEW

Appendix 1.2. 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 Atlantic 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, ○ Coral species occurring in region 

but not identified by regional author as major structure-forming species. Deep-water corals reported by a 

circle under “Caribbean” heading are from the U.S. Caribbean only. ~ Indicate strcuture forming coral found in 

Caribbean but not in U.S. waters. 

Higher Taxon Species Northeast Southeast Gulf of Mexico Caribbean

Phylum Cnidaria

Class Anthozoa

Subclass Hexacorallia 

Order Scleractinia

Family Caryophylliidae Lophelia pertusa ○ ● ● ●

Solenosmilia 

variabilis ○ ○ ● ~

Desmophyllum 

dianthus ○ ○ ○ ●

Family Dendrophylliidae

Enallopsammia 

profunda ○ ● ● ~

Enallopsammia 

rostrata ○ ○ ●

Dendrophyllia 

alternata ○ ~

Family Oculinidae Madrepora oculata ● ● ●

Madrepora carolina ○ ● ●

Oculina varicosa ● ○ ~

Family Pocilloporidae Madracis myriaster ● ○ ●

Order Antipatharia

Family Antipathidae

Antipathes 

americana ●

Antipathes 

caribbeana ●

Family Leiopathidae

Leiopathes 

glaberrima ● ● ~

Family Myriopathidae

Plumapathes 

pennacea

○ ●

Tanacetipathes hirta ●

Family Schizopathidae

Bathypathes 

alternata ● ○ ~

Parantipathes 

tetrasticha ●

Subclass Octocorallia 

Order Gorgonacea

Family Acanthogorgiidae

Acanthogorgia 

armata ● ~
STATE OF DEEP CORAL ECOSYSTEMS OF THE UNITED STATES

INTRODUCTION AND OVERVIEW

64

Higher Taxon Species Northeast Southeast Gulf of Mexico Caribean

Family Anthothelidae

Diodogorgia 

nodulifera ○ ●

Family Ellisellidae

Ellisella 

barbadensis ○ ●

Ellisella elongata ○ ●

Nicella 

deichmannae ●

Nicella 

guadelupensis ○ ●

Nicella obesa ●

Riisea paniculata ○ ●

Family Isididae Acanella arbuscula ○ ●

Keratoisis flexibilis ○ ● ~

Keratoisis spp. ○ ●

Family Paragorgiidae Paragorgia arborea ●

Family Plexauridae

Paramuricea 

grandis ● ~

Swiftia exserta ●

Family Primnoidae

Acanthoprimnoa 

goesi ●

Callogorgia 

americana 

americana

○ ○ ●

Callogorgia 

americana delta ●

Narella bellissima ○ ●

Narella pauciflora ○ ●

Primnoa 

resedaeformis ●
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

65

Auke Bay Laboratory, Alaska Fisheries Science 

Center

National Marine Fisheries Service

11305 Glacier Highway

Juneau, Alaska 99801-8626

STATE OF DEEP CORAL ECOSYSTEMS

IN THE ALASKA REGION:

GULF OF ALASKA, BERING SEA AND THE ALEUTIAN ISLANDS

Robert P. Stone and S. Kalei Shotwell

I. INTRODUCTION

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 

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 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

66

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 II. 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 

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).
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

67

Alexander Archipelago, Southeast Alaska, 2) the 

western GOA including the Alaska Peninsula, 3) 

the eastern Aleutian Islands (Shumagin Islands 

to Seguam Pass), 4) the western Aleutian Islands 

(Seguam Pass to Stalemate Bank), and 5) the 

Bering Sea. 

Coral records from these areas were categorized 

into the six major taxonomic groups. Three 

ecologically important groups of gorgonians, 

Primnoa spp., Paragorgia spp., and bamboo 

corals (Family Isididae) are categorized separately 

because their large size and conspicuous 

morphology greatly reduce the probability of 

inaccurate field identification. 

The principal source of information on coral 

distribution is by-catch data collected during NMFS 

research trawl surveys (Resource Assessment 

and Conservation Engineering Database 

(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). 

II. GEOLOGICAL SETTING

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, 

and methane seeps (NURP 2001). The nature 

of the seabed on the Gulf of Alaska shelf has 

been strongly influenced by glaciation and high 

rates of sediment deposition. The Gulf of Alaska 

also contains approximately 24 major seamounts 

arranged in three chains extending north from 

the Juan de Fuca Ridge. The seamounts are 

volcanoes rising from the abyssal plain that were 

likely formed as the Pacific Plate moved over 

mantle hotspots. 

The Bering Sea

The Bering Sea is a shallow sea and has one of 

the largest continental shelves in the world – 1200 

km long and 500 km wide (National Academy of 

Sciences 1990). The continental shelf breaks 

at approximately 170 m depth and seven major 

canyons, including the Zhemchug and Bering 

Canyons—the two largest submarine canyons 

in the world (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. The Aleutian Island Archipelago 

contains more than 300 islands and extends 

over 1900 km from the Alaska Peninsula to the 

Kamchatka Peninsula in Russia. The Archipelago 

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 longshore currents, wave action, entrainment in 

bottom-fast ice, ice gouging, ocean currents, and 

internal waves (Horowitz 2002). 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

68

III. OCEANOGRAPHIC SETTING

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 North America, 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 

2003).

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 

eastward. 

IV. STRUCTURE AND HABITAT-FORMING 

DEEP CORALS

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 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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69

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 

Scleractinia)

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 structureforming corals (Table 2.1) and at least 14 species 

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.

Taxa

Reefbuilding AbundanceMaximum 

colony size Morphology

Associations 

with other 

structure 

forming 

invertebrates

Colony 

spatial 

dispersion

Overall 

rating of 

structural 

importance

Scleractinia No Low Small No-branch Few Clumped Low

Antipatharia No Medium Large Branch Few Clumped Medium

Alcyonacea No Medium Small No-branch Few Clumped Medium

Stolonifera No Low Small No-branch Few Clumped Low

Gorgonacea No High Large Branch Many Clumped High

 Primnoa spp. No High Large Branch Many Clumped High

Paragorgia

spp. No Medium Large Branch Many Clumped High

Isididae No Medium Large Branch Many Clumped High

Pennatulacea No High Large Branch Few Clumped Medium

Anthoathecatae No High Medium Branch Many Clumped High

Table Key

Attribute Measure

Reef-Building Yes/No

Relative Abundance Low/ Medium/ High

Size (width or height) Small (<30cm)/ Medium (30cm-1m)/ Large (>1m)

Morphology Branching/ Non-branching

Associations None/ Few (1-2)/ Many (>2)

Spatial Dispersion Solitary/ Clumped

Overall Rating Low/ Medium/ High

Table 2.1. Structure-forming attributes of deep corals in Alaska.

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 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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70

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.

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

 Zoanthidae) 

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) 

indicate that the 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 singleand 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 of Alaska (Krieger 2001) 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 3.7oC (Cimberg et al. 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 2.3. Some black corals such as this Dendrobathypathes boutillieri may reach heights over 1 m. 

An unknown species of octopus takes cover under the 

coral. Photo credit: R. Stone, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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71

a temperature tolerance of less than 3.0oC 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 

 Pennatulacea)

Ten species of pennatulaceans (sea pens) are 

reported from Alaskan waters (Appendix 2.1) 

and several are important structure-forming 

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 structureforming 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 structureforming invertebrates.

Figure 2.5. A large bubblegum coral (Paragorgia arborea) provides shelter for a Pacific cod 

(Gadus macrocephalus) in the central Aleutian Islands. Photo credit: R. Stone, NOAA Fisheries.

Figure 2.6. 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 Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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72

V. SPATIAL DISTRIBUTION OF CORAL 

SPECIES AND HABITAT

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 

subspecies of stylasterids (Appendix 2.1). 

Corals range in depth from 6 m for Primnoa pacifica

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 (Krieger 2001). 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 bottomcontact 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 

Figure 2.7. Large, erect stylasterids (Stylaster sp). 

grow on exposed bedrock with their central axis perpendicular to the current in the Aleutian Islands. Red 

laser marks are separated by 10 cm. Photo credit: 

R. Stone, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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73

Figure 2.8. Distribution of corals in the eastern Gulf of Alaska A) gorgonians (bamboo corals – Family Isididae, Paragorgia spp., Primnoa spp. are plotted separately), B) stylasterids, C) black corals, D) stony corals, E) 

soft corals, and F) pennatulaceans.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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74

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 of Alaska 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 of Alaska (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 most abundant 

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. 

comm.).

Coral habitat on Derickson 

Seamount which crests at 

2766 m south of the Alaska 

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! Deepwater 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, 

Taxa

Number of 

Species Depth range (m)

Data source 

shallow - deep

Scleractinia 11 24 - 4620 4 - 3

Antipatharia 14 401 - 4784 4 - 1

Alcyonacea 9 10 - 3209 4 - 2

Stolonifera 6 11 - 591 6 - 4

Gorgonacea 63 6 - 4784 5 - 1

Pennatulacea 10 3 - 2947 4 - 4

Anthoathecatae 28 11 - 2130 6 - 4

Total 141 3 - 4784

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.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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75

Figure 2.9. Distribution of corals in the western Gulf of Alaska A) gorgonians (bamboo corals – Family Isididae, Paragorgia spp., Primnoa spp. are plotted separately), B) stylasterids, C) black corals, D) stony corals, E) 

soft corals, and F) pennatulaceans.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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76

unpublished data) including Dendrobathypathes 

boutillieri, a species new to science (Opresko 

2005).

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 

169o W near the 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 report we 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 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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77

Figure 2.10. Distribution of corals in the eastern Aleutian Islands A) gorgonians (bamboo corals – Family Isididae, Paragorgia spp., Primnoa spp. are plotted separately), B) stylasterids, C) black corals, D) stony corals, E) 

soft corals, and F) pennatulaceans.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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78

Figure 2.11. Distribution of corals in the western Aleutian Islands A) gorgonians (bamboo corals – Family Isididae, Paragorgia spp., Primnoa spp. are plotted separately), B) stylasterids, C) black corals, D) stony corals, E) 

soft corals, and F) pennatulaceans.
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79

Figure 2.12. Distribution of corals in the Bering Sea A) gorgonians (bamboo corals – Family Isididae, Paragorgia spp., Primnoa spp. are plotted separately), B) stylasterids, C) black corals, D) stony corals, E) soft corals, 

and F) pennatulaceans.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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80

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 71o 

24’ N. 

VI. SPECIES ASSOCIATIONS WITH DEEP

CORAL COMMUNITIES 

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 

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 

Figure 2.13. A shortspine thornyhead (Sebastolobus 

alascanus) rests in a field of primnoid gorgonians. 

Photo credit: R. Stone, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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81

physical contact with corals and other structureforming 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), 

and several species of grenadiers (Family 

Macrouridae). 

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. For example, three species of Aleutian 

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). 

All megafauna were in physical contact 

with the coral and were using it as an 

elevated feeding platform or as refuge. 

The spiny red sea star (Hippasteria 

spinosa) was documented preying on 

the coral. Macrofauna such as shrimp 

were also observed within the colonies 

but were not identified or enumerated. 

Figure 2.15. An unidentified eelpout (probably Puzanovia rubra) displays cryptic coloration in a Paragorgia colony at 746 m depth in the Aleutian Islands. 

Photo credit: R. Stone, NOAA Fisheries.

Figure 2.14. A darkfin sculpin Malacocottus zonurus rests under 

a bubblegum coral Paragorgia arborea in one of the seven coral 

gardens surveyed with the submersible Delta. Coral gardens 

are areas of extraordinary coral abundance and high species 

diversity. Photo credit: R. Stone, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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82

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 appear to 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 

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.

VII. STRESSORS ON DEEP CORAL 

COMMUNITIES 

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 

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 discovered and are currently being described by Dr. James 

McLean at the Natural History Museum of Los Angeles County. Photo credit: R. Stone, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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83

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 bycatch, 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 central Aleutian 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 

2006). 

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 widescale ecosystem change (Gislason 1994; Goñi 

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 142o 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 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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84

Figure 2.17. Average 

annual bottom trawl fishing 

intensity between 1998 

and 2002 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 Rose 

and Jorgensen 2005). 

Trawl intensity is defined in 

NMFS (2005), Appendix B. 

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 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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85

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 

2.3).

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.

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 

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 

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 

Gear Type

Severity 

of effects

Extent of 

effects

Geographic 

extent of use

Overlap of 

use with 

coral habitat

Overall rating 

of gear 

effects

Otter trawls High High High Medium High

Mid-water trawls Low Low Medium Low Low

Demersal longlines Medium Low High Medium Medium

Single-set pots Low Medium Medium Low Low

Longline pots High Medium Low Medium Medium

Scallop dredges Medium Low Low Low Low

Table 2.3. Potential effects of fishing gears on deep coral habitat in Alaska.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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86

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 

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). Methodology described in Fritz et 

al. 1998.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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87

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 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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88

Figure 2.20. Number of observed crab pots during the golden king crab (Lithodes aequispina) fishery between 2001 and 2004 in A) the eastern Aleutian Islands and B) the western Aleutian Islands. Each area rectangle measures 0.05o latitude by 0.1o longitude. Source data: Alaska Department of Fish and Game.

coral habitat since they generally occur in softsediment areas with minimal coral habitat and 

because a relatively small area of the seafloor is 

contacted with the gear (Table 2.3). Pot longlines 

used in the Aleutian Island golden king crab 

fishery, however, have the potential to cause 

extensive damage to coral habitat (Table 2.3) 

since the spatial distribution of fishing is extensive 

in some areas of high coral abundance (Figures 

2.20A and 2.20B). Depending on how the gear 

is retrieved, the area of seafloor contacted may 

be relatively large and the forces on the seafloor 

may be substantial. The gear is retrieved in a 

manner to minimize drag on the seafloor due to 

the strength limitations of the longline; however, 

under certain conditions the gear can be dragged 

like a plough across the seafloor. This situation 

can occur in areas of steep bathymetry and when 

strong winds and currents dictate that fishing 

vessels retrieve gear while being forced away 

from it. At one site in the central Aleutian Islands 

where disturbance from this gear was observed 

with the submersible Delta (Figure 2.21), the 

seafloor was scoured to bare substrate along 17 

strips (Stone 2006). Aleutian Island coral gardens 

are at high risk to disturbance from this fishery.

Scallop Dredges

A small but important fishery has occurred for 

the weathervane scallop Patinopecten caurinus 

in the Gulf of Alaska and Bering Sea since 1967 

(Shirley and Kruse 1995). The fishery occurs in 

relatively well-defined areas of unconsolidated 

soft sediments on the continental shelf and 

at depths between 60 and 140 m (Turk 2001; 

Barnhart 2003). Scallop dredges are dragged 

along the seafloor and designed to dig into the 

top layer of sediment. Dredges have a maximum 

width of 4.6 m. No directed research on the 

effects of scallop dredges on coral habitat has 

been undertaken in Alaska. Overlap does occur 

between the fishery and the known distribution of 

pennatulaceans, including ecologically important 

groves of Halipteris willemoesi in the central 

Gulf of Alaska (Masuda and Stone 2003). With 

the exception of pennatulacean groves, scallop 

dredges pose a minimal threat to coral habitat 

at the present time (Table 2.3). The spatial 

Figure 2.21. Coral habitat can be fragmented by 

fishing gear as evidenced by this small patch of 

coral in the path of heavy disturbance. A small 

bubblegum coral (Paragorgia arborea), soft coral 

(Anthomastus sp.), stoloniferan coral (Clavularia sp.), hydrocorals, demosponges, and a sea 

anemone reside at the edge of a path littered with 

hydrocoral skeletons. Photo credit: R. Stone, 

NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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89

distribution of fishing is small and the area of the 

seafloor contacted per tow is relatively small. 

While the gear is intrusive, it is generally used 

in soft-sediment areas where coral abundance 

is low. Groves of pennatulaceans in the Gulf 

of Alaska are most at risk to disturbance from 

scallop dredges. 

Effects of Other Human Activities

Oil and Gas Exploration and Extraction

Offshore oil and gas operations in Alaska include 

exploration, development, and production 

activities (NMFS 2005). Most of these activities 

presently occur in Cook Inlet in the Gulf of Alaska 

and on the North Slope (Beaufort and Chukchi 

Seas)—areas of Alaska that do not support 

significant deep coral communities. Disturbances 

from these activities that may affect coral habitat 

include physical alterations to habitat, waste 

discharges (well drilling cuttings and muds), 

and oil spills. Cimberg et al. (1981) discuss 

the potential effects of oil and gas development 

on deep corals in Alaska. They concluded that 

Alaskan corals are unlikely to suffer adverse 

effects from oil and gas development, because 

most of the known deep coral distributions do not 

occur in lease areas and areas where platforms 

will be placed. They further conclude that the 

most physiologically sensitive life history stage of 

deep corals, the planula larval stage, is brief and 

demersal, and therefore unlikely to be affected 

by oil spills (Cimberg et al. 1981). No directed 

research has been undertaken to study the effects 

of oil toxicity on any life history stage of deep 

corals found in Alaska. The potential for effects 

from these activities on coral habitat is likely to 

increase in the future as the world’s demand for 

oil and gas products continues to increase. 

Deployment Of Petroleum Pipelines And 

Communication Cables

Cook Inlet is the only area of the state where 

petroleum pipelines (specifically crude oil) have 

been deployed in benthic marine habitats. 

Fourteen pipelines totaling 141 km in length were 

deployed on the seafloor of Cook Inlet between 

1965 and 1986 (Robertson 2000). Eight state and 

federal agencies have regulatory authority over 

pipelines in Cook Inlet. Accidental spills have 

occurred in the past and are likely to occur in the 

future as many of the pipelines reach the end of 

their expected life span (Robertson 2000). The 

benthic marine life in the immediate area of the 

pipelines has not been inventoried but the region 

is not known to support abundant or diverse coral 

resources. 

Numerous communication cables have been 

deployed on the ocean floor throughout Alaskan 

waters since 1900. Thousands of kilometers 

of cables stretch along the seafloor between 

Alaskan communities and ports in Washington 

and Oregon. Cables have been deployed 

from the shoreline down to depths of 7000 

m. There are no known regulations governing 

the placement of submarine cables but their 

locations are accurately mapped so that potential 

interactions with other seafloor uses (e.g., fishing) 

can be avoided. There are no known reports of 

cable deployments directly affecting coral habitat 

although there is some likelihood that cables 

have been placed in coral habitat, especially in 

the Aleutian Islands. Cables are typically laid 

on the seafloor where they remain exposed but 

may be buried using specially designed ploughs 

in areas where bottom fishing and other seafloor 

uses occur. In areas where cables are exposed 

they may provide attachment substrates for 

corals and other emergent epifauna and may 

therefore provide a known time-line for studies of 

recruitment and subsequent growth of emergent 

epifauna that settle on them (Levings and 

McDaniel 1974). No such studies have been 

conducted yet in Alaska but clearly the potential 

exists to use submarine cable deployments to 

gain insights into coral habitat recovery rates. 

Deployment of communication cables is presently 

a minimal threat to deep coral ecosystems in 

Alaska given the very small area of the seafloor 

that is contacted by them. 

Pollution - Point-source Discharges

Point-source discharges that occur in coastal 

marine areas of Alaska have little potential to affect 

deep coral habitat. Coral habitat is sparse in coastal 

areas of Alaska where point-source discharges 

occur or are expected to occur in the near future 

but a few coastal areas near municipalities 

do support groves of pennatulaceans. The 

greatest threat to coral habitat from point-source 

discharges is the introduction of large volumes 

of untreated sewage and chlorine. Sewage 

discharge causes organic nutrient enrichment 

of receiving waters, rapid increases in biological 

production, and eutrophication through depletion 

of dissolved oxygen (Tomascik et al. 1997). High 

concentrations of phosphates in effluent may 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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90

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 either singleevent acute exposures or to chronic exposures 

(Tomascik et al. 1997).

Fish Processing Waste

In Alaska, 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 bycatch, 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 Niño/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 CO2 emissions 

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 structureforming 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 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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91

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.

VIII. MANAGEMENT OF FISHERY 

RESOURCES AND HABITATS

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 

habitat. 

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 et al. 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 seafloor with 

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://

www.nprb.org/research/res_2003_projects_

listing.htm) is to develop a model to predict 

where deep coral habitat is located throughout 

the region. Mapped areas were systematically 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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Region Name of area Agency

Depth range 

(m)

Coral 

presence Area (km2)

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

GOA 

Albatross Bank, 

Snakehead NMFS 60-810 Yes 310

GOA 

Albatross Bank, 8-fathom 

pinnacle NMFS, NOS 20-716 No 17

GOA 

Albatross Bank, 49-

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

GOA, SM

Transit between 

seamounts UAF, OSU ? Unknown 9,000

GOA, SM Derickson Seamount UAF, OSU 2750-6800 Yes 5,200

AI Akutan UAF 78-482 Unknown 27

AI Bogoslof UAF 20-820 Unknown 28

AI RV Revelle transit UAF 90-4200 Yes 11,341

AI Samalga Island, North NMFS 107-323 Yes 11

AI Samalga Island, South NMFS 120-150 Yes 9

AI

Islands of Four 

Mountains, North NMFS 144-223 Yes 13

AI

Islands of Four 

Mountains, South NMFS 88-204 Yes 12

AI

Islands of Four 

Mountains, West NMFS 116-218 Yes 11

AI Aleutian Corals NMFS 100-3000 Yes 2,697

AI Track Lines NMFS 30-4000 Yes NA

Total 46,710

Table 2.4. Areas of Alaska that have been mapped with modern multibeam sonar technology. AI = 

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.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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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 

features.

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 

causes. 

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, Southeast Alaska 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 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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comprehensive inventory and classification of 

Marine Protected Areas (MPAs) in Alaska and a 

brief history of their development is provided in 

Witherell and Woodby (2005).

In February 2005 the NPFMC voted unanimously 

to protect vast areas of seafloor habitat in the Gulf 

of Alaska and Aleutian Islands. NMFS approved 

the Record of Decision on the NPFMC essential 

fish habitat (EFH) environmental impact statement 

(EIS) containing these provisions on August 8, 

2005 and the regulations were implemented in 

July 2006. The Aleutian Islands closures include 

a vast area (957, 361 km2) of seafloor west of the 

Islands of Four Mountains (170° W) and includes 

nearly the entire EEZ in the region (Figure 2.24). 

The Aleutian Islands Habitat Conservation Area 

(AIHCA) is the largest bottom trawl closure in the 

U.S. and the first in the state directed at protecting 

sensitive deep coral habitat. Under the decision, 

areas that have been trawled in the past and 

have supported the highest groundfish catches 

will remain open (42,611 km2), while largely 

unfished areas including Bowers Ridge in the 

Bering Sea, are closed to bottom trawling (Figure 

2.24). Approximately 52% of the fishing grounds 

previously targeted by trawlers (defined as those 

habitats within the current depth range of trawl 

activities—1000 m depth), or about 100,000 km2 

of seafloor, are now closed to bottom trawling in 

the Aleutian Islands (Figure 2.24). The majority of 

the seafloor habitat within the closure has not yet 

been explored by scientists but much of it is deep 

abyssal plain that likely supports little deep coral 

habitat. Initial indications are that few species of 

commercial importance exist in these deep areas 

that would have put the habitat at risk to immediate 

or future trawling activities. Nonetheless, the 

closure effectively freezes the current footprint of 

trawling activities until scientists can determine 

the full scope of deep coral habitat and fisheries 

resources in the region. Perhaps the highlight 

Figure 2.22. Areas of Alaska with some restrictions on fishing activities. Map credit: Cathy Coon, NPFMC.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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of the AIHCA is the closure of six coral gardens 

(377 km2 total) in the central Aleutian Islands 

where the use of all bottom-contact gear is now 

prohibited (Figure 2.24). 

The NPFMC also voted to establish HAPCs in the 

Gulf of Alaska where the use of all bottom-contact 

fishing gear would be prohibited. These include 

five small areas (46 km2 total) in the eastern Gulf 

of Alaska to protect dense thickets of red tree 

corals (P. pacifica), 7,155 km2 of slope habitat 

and 16 seamounts (the majority in the Gulf of 

Alaska) to protect deep coral habitat (18,278 km2 

total). 

the State of Alaska to regulate fishing vessels 

in the EEZ, even if not registered with the State, 

if they are operating in a fishery for which there 

was no FMP in place on August 1, 1996. In 2003, 

the State of Alaska adopted a regulation (5 ACC 

38.062e) that states that no permits will be issued 

to commercially harvest corals or sponges. This 

regulation applies to both state waters and those 

within the EEZ. 

Minerals Management Service

Offshore drilling activities are regulated by 

the U. S. Department of the Interior’s Minerals 

Management Service and their proposed plan for 

outer continental shelf (OCS) oil and gas leasing 

Figure 2.23. Primnoa pacifica thrives in the cold, dark glacial fjords of Southeast Alaska. 

This colony, at a depth of only 18 m, provides a resting platform for a blue king crab (Paralithodes platypus). Photo credit: R. Stone, NOAA Fisheries.

Directed Harvest

In April 2000, the NPFMC introduced a regulation 

defining all corals as prohibited species (Witherell 

and Coon 2001). The regulation would have 

effectively prohibited the sale, barter or trade of 

corals but the measure was never adopted by 

the NPFMC since it would only apply to federally 

licensed groundfish vessels and fishermen. 

Rather than adopt a measure that would not 

have applied to all fishing vessels, the NPFMC 

relegated management authority to the State of 

Alaska under provisions of the MSFCMA. Section 

306(a)(3)(C) of the MSFCMA gives authority to 

(2007 - 2012) includes two lease sales in the 

North Aleutian Basin. Offshore drilling in the OCS 

region of Alaska began in 1975 and since that time 

nearly 100 exploratory wells have been drilled. 

Twenty nine wells have been drilled in shallowwater (5 - 51 m depth) areas of the Beaufort Sea. 

Five wells have been drilled at depths between 

42 and 46 m in the Chukchi Sea, 24 wells have 

been drilled in the Bering Sea Region (Norton 

Sound, Navarin Basin, and south of the Pribilof 

Islands) at depths between 11 and 165 m, and 

25 wells have been drilled in the Gulf of Alaska 

including 13 in Cook Inlet at depths between 35 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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96

and 263 m. There are currently no regulations 

in place specifically to protect deep corals from 

these activities and most present drilling activities 

occur in areas not known to support abundant or 

diverse coral resources. In early 2007 however, 

the longstanding moratorium barring offshore oil 

and gas drilling in Bristol Bay and the southeastern 

Bering Sea was lifted by President Bush paving 

the way for oil and gas lease sales in this region 

of Alaska. While removal of the ban has caused 

the greatest concern over the potential effects 

offshore drilling may have on the area’s rich 

stocks of salmon and groundfish, the area may 

also support important coral habitat, particularly 

beds of soft corals, that could be affected by any 

development in the region. 

National Marine Sanctuaries

No National Marine Sanctuaries exist in Alaskan 

waters.

IX. REGIONAL PRIORITIES TO 

MANAGE AND CONSERVE DEEP 

CORAL COMMUNITIES 

Directed research on deep coral habitat has 

been undertaken only during the past decade 

in the Alaska Region. Geographical areas 

known to support abundant and diverse deep 

corals, such as the Aleutian Islands and Gulf of 

Alaska seamounts, have been the first priority 

for exploration and specimen collection. While 

specimen collections and direct observations of 

deep corals in those areas will provide important 

foundation studies on coral ecology and 

systematics, many research priorities remain for 

the region.

Recently, NMFS recommended that the NPFMC 

pursue three courses of action regarding the 

effects of fishing on essential fish habitat in 

Alaska (NMFS 2005) and these apply directly to 

deep coral habitat:

1. The NPFMC and NMFS should continue 

to analyze how implementation of fisheries 

management measures affects seafloor 

habitats.

2. The NPFMC should continue to support 

research to improve scientific understanding 

of the effects of fishing on seafloor habitat, 

the ecological processes linking habitats and 

managed species, and the recovery rates of 

seafloor habitats disturbed by fishing gear.

3. The NPFMC should take specific 

precautionary management actions to avoid 

Figure 2.24. Closure areas in the Aleutian Islands, Alaska that were proposed by the North 

Pacific Fishery Management Council in February 2005 and approved by NOAA Fisheries in 2006. 

Map credt: Cathy Coon, NPFMC.
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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additional disturbance to fragile seafloor 

habitats that may be especially slow to 

recover.

The third course of action recommended by 

NMFS has already been met to a large degree 

through the recent closure of vast areas of 

seafloor to fishing activities. These closures 

now provide an excellent opportunity to pursue 

the first and second courses of action to some 

degree. In support of these courses of action we 

have identified the following priorities for future 

deep coral research and conservation in the 

Alaska Region. 

• Research

Studies on the growth rates and reproductive 

ecology from representative corals of the major 

taxonomic groups need to be undertaken to provide 

estimates of recovery rates for coral habitat. 

These data will provide a better understanding of 

the ability of species to recover from disturbance 

and recolonize areas set aside as mitigative 

measures such as HAPCs or MPAs. Growth rates 

are known for only three octocoral species that 

occur in Alaska (Calcigorgia spiculifera, Halipteris 

willemoesi, and Primnoa sp.) and range from 

approximately 5.8 mm yr-1 to 23.2 mm yr-1 (Stone 

and Wing 2002; Wilson et al. 2002; Andrews et 

al. 2002). Whether these rates encompass the 

full range of growth rates for other octocorals or 

for other taxonomic groups (e.g., antipatharians) 

is unknown. Additional growth rate studies are 

needed for representative species from each 

major structure forming taxonomic group and for 

those taxa that form single-species assemblages 

that provide important habitat in certain regions. 

The following taxa are excellent candidates 

for growth studies: 1) the gorgonians Fanellia

spp., Plumarella spp., Thouarella spp., and 

Keratoisis spp. or Isidella spp., 2) the stylasterids 

Stylaster cancellatus and S. campylecus, 3) the 

antipatharians Chrysopathes formosa and C. 

speciosa, and 4) the true soft coral Eunephthea 

rubiformis. Studies on the reproductive ecology 

of deep corals have been limited to Aleutian 

Island stylasterids (Brooke and Stone in press) 

and a few species of Aleutian Island gorgonians 

(Anne Simpson, University of Maine, unpublished 

data). Reproductive ecology studies should be 

undertaken for Primnoa spp. and all of the taxa 

listed above. 

Taxonomic studies need to be expanded so 

that accurate identifications of by-catch can be 

made in the field. Gorgonians, stylasterids, black 

corals, pennatulaceans, and true soft corals are 

abundant and important structure-forming corals 

in the Alaskan Region yet their taxonomy is still 

poorly understood despite the fact that extensive 

collections have now been made throughout 

much of the region. The taxonomy for 52 of 

the 141 unique taxa of deep corals (nearly 

37%) documented from Alaska have not been 

accurately identified to species. 

Studies on the effects of ocean acidification and 

oil toxicity on deep corals should be undertaken 

now. Ocean acidification could have serious 

consequences on deep corals in areas of the 

North Pacific Ocean where the aragonite and 

calcite saturation horizons are already quite 

shallow and predicted to become shallower in the 

near future (Guinotte et al. 2006). Studies should 

include monitoring shallow-water populations of 

the scleractinians Javania borealis, Caryophyllia 

alaskensis, and Balanophyllia elegans in the 

central Aleutian Islands (Stone 2006) and 

also laboratory manipulative experiments that 

subject cold water corals to the more acidic 

conditions they are likely to encounter in the 

coming decades. New deep-sea technology 

(e.g. free ocean CO2 enrichment systems or 

FOCE systems) is currently being developed 

that will allow scientists to monitor calcification 

rates of corals in situ while carefully controlling 

and manipulating pH (Bill Kirkwood, Monterey 

Bay Aquarium Research Institute, pers. comm). 

When fully developed this technology will be a 

highly valuable tool for studying the effects of 

ocean acidification on deep corals in Alaska. 

Oil and gas exploration and extraction is likely 

to increase in the marine environment of Alaska 

in the near future. Laboratory manipulative 

experiments should be conducted to determine 

the toxicity effects of oil on both the adult and 

larval stages of deep corals. The soft coral, 

Eunephthea rubiformis, is an excellent candidate 

for such studies. 

Studies on the effects of specific fishing gear 

types on coral habitat need to be undertaken 

so that we can better understand the effects of 

certain fisheries on coral habitat. Experiments on 

gear modifications to reduce coral by-catch and 

contact with the seafloor should be a high priority. 

Further studies need to be conducted on the use 

of coral habitat by managed groundfish species. 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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Limited research to date clearly indicates that 

many FMP species are associated with coral 

habitat but the nature of the associations is still 

unknown. Additional research is necessary to 

examine in detail how these species use coral 

habitat (i.e., to accomplish which life processes) 

so that estimates of changes to overall fisheries 

productivity can be made following disturbance.

Reconnaissance submersible dives should be 

undertaken in coral “hotspots” to assess their 

suitability as HAPCs. Coral hotspots include 

areas of abundant and diverse by-catch and 

areas known or suspected by scientists to support 

assemblages of gorgonians, pennatulaceans, 

and antipatharians. Hotspots include areas 

along the shelf and upper slope of the Gulf of 

Alaska where stock assessment surveys indicate 

the presence of Primnoa thickets and patches of 

bamboo corals and black corals. Other hotspots 

include several of the canyons on the Bering Sea 

shelf where survey by-catch records indicate 

the presence of Primnoa sp, Paragorgia sp., 

bamboo corals, and groves of pennatulaceans—

the northernmost records for these corals in the 

North Pacific Ocean.

Reconnaissance submersible dives should also 

be made on areas of Bowers Bank in the Bering 

Sea that were recently closed to all bottom 

trawling as part of the recently established 

Aleutian Islands Habitat Conservation Area. The 

Bank is completely unexplored and is one of the 

few regions of Alaska that has received very little 

fishing pressure but is thought to contain extensive 

deep coral habitat based on its geographical 

proximity to the Aleutian Islands and Petrel Bank 

where rich coral habitat has been documented.

• Seafloor mapping

Multibeam mapping technology is very expensive 

(i.e., recent efforts in the Aleutian Islands were 

approximately $30,000 per day and on average 

about 80 km2 could be mapped per day) and 

consequently only a fraction of 1% of the seafloor 

in the Alaska Region has been mapped to date. 

Some of the recent mapping effort has been 

in support of submersible operations with the 

directed coral studies. Other efforts have been 

in heavily fished areas in the Gulf of Alaska that 

do not support abundant coral resources but 

are important areas for the study of gear effects 

on seafloor habitat. Many areas of high coral 

abundance, based on by-catch records, have 

not been mapped and they should rate as a high 

priority for the limited funds available for that 

purpose. The coral hotspots listed above are a 

high priority for mapping. 

• Coral by-catch database

Coral by-catch in fisheries and stock assessment 

surveys will continue to be an important data 

source in mapping Alaska’s coral resources and 

interactions with fisheries. Databases for these 

surveys are well established and maintained and 

are a tremendous source of data (e.g., Heifetz 

2002; Etnoyer and Morgan 2005). While the 

databases are currently in a usable format some 

restrictions regarding confidentiality do limit their 

usefulness at the present time. Fisheries observer 

databases could be greatly improved by including 

more precise locations of fishing effort and coral 

by-catch. Coral by-catch from the surveys is also 

a tremendous, and largely untapped, source of 

specimens for taxonomic, paleoclimatological, 

and ecological analyses. Specific collection 

requests for these purposes are occasionally 

accepted by the survey teams but little time 

is typically available for these purposes and 

proper identification of requested taxa are often 

difficult. Training in coral identification should 

be made available to all fisheries observers and 

the publication of additional field identification 

guides should be undertaken to facilitate these 

requests. 

X. CONCLUSION

Deep corals are an important component of 

benthic ecosystems in Alaska. Highly varied 

submarine geology, persistent water currents, and 

plankton rich waters support at least 141 species 

from six major taxonomic groups. Deep corals 

have a broad geographical and depth distribution 

within the region. The Aleutian Islands support 

the highest diversity and abundance of corals in 

Alaskan waters but other subregions, such as the 

Gulf of Alaska and Bering Sea, support important 

single-species assemblages of gorgonians, 

pennatulaceans, and true soft corals. Many are 

important structure-forming species due to their 

large size, branching morphology, and patterns 

of distribution. 

Fishing gear that contacts the seafloor is 

presently the single largest threat to coral habitat 

in Alaska. Evidence from research submersible 
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

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observations and coral by-catch data collected 

during fisheries and stock assessment surveys 

indicate that interactions between coral habitat 

and current fisheries occur in many areas. 

The NPFMC and NMFS have taken significant 

steps to address this interaction by recently 

implementing extensive closures specifically to 

protect coral habitat from disturbance caused by 

fishing activities.

Most of the commercial species currently 

harvested in Alaska (approximately 85% of all 

FMP species or species groups) spend all or part 

of their life cycle in deep habitat where corals 

are potentially found. As the world population 

continues to grow and the demand for seafood 

increases in the future, conservation of Alaska’s 

deep coral resources will be a major challenge 

for managers striving to maintain sustainable 

fisheries. NOAA recognizes the value of both 

shallow and deep coral habitat conservation 

in that endeavor and has listed it as one of 

nine programs within the Ecosystems goal 

in its strategic plan—the only taxa explicitly 

listed in the Strategic Plan and the MagnusonStevens Fishery Conservation and Management 

Reauthorization Act of 2006 mandates continued 

research, mapping, and protection of deep coral 

communities. 

XI. REFERENCES

Aagaard K (1984) The Beaufort Undercurrent. 

In: Aagaard K, Barnes PW, Schell DM, 

Reimnitz E (eds). The Alaskan Beaufort 

Sea: ecosystems and environments. 

Academic Press, Orlando, Florida. 466 

pp

ADF&G (Alaska Department of Fish and Game) 

(2002) Alaska Aquatic Nuisance Species 

Management Plan. Alaska Department 

of Fish and Game, Commissioner’s 

Office, Juneau, Alaska

Alcolado PM, Claro R, Menendez G, GarciaParrado P, Martínez-Daranas B (1998) 

Status of Cuban coral reefs. Atlantic Atlantic 

and Gulf Reef Assessment Workshop. 

June 2-6, 1998. Available at Institute 

of Oceanography, Ministry of Science, 

Havana, Cuba

Allen JS, Beardsley RC, Blanton JO, Boicourt 

WC, Butman B, Coachman LK, Huyer 

A, Kinder TH, Royer TC, Schumacher 

JD, Smith RL, Sturges W, Winant CD 

(1983) Physical oceanography of 

continental shelves. U.S. National 

Report to International Union of 

Geodesy and Geophysics 1979-1982. 

Reviews of Geophysics and Space 

Physics 21:1149-1181

Alton SM (1986) Fish and crab populations of 

the Gulf of Alaska seamounts. In: 

Uchida RN, Hayasi S, Boehlet GW 

(eds). Environment and resources of 

seamounts in the North Pacific. U.S. 

Department of Commerce, NOAA 

Technical Report NMFS 43.Anderson 

ME (1994) Systematics and osteology of 

the Zoarcidae (Teleostei: Perciformes). 

Ichthyological Bulletin of the J. L. B. 

Smith Institute of Ichthyology 60:120 

pp

Andrews AH, Cordes EE, Mahoney MM, Munk 

K, Coale KH, Cailliet GM, Heifetz J 

(2002) Age, growth and radiometric age 

validation of a deep-sea, habitat-forming 

gorgonian (Primnoa resedaeformis) 

from the Gulf of Alaska. Hydrobiologia 

471:101-110

Auster PJ, Malatesta RJ, Langton RW, Watling 

L, Valentine PC, Donaldson CLS, 

Langton EW, Shepard AN, Babb IG 

(1996) The impacts of mobile fishing 

gear on seafloor habitats in the Gulf of 

Maine (northwest Atlantic): implications 

for conservation of fish populations. 

Reviews in Fisheries Science 4:85-202

Auster PJ, Langton RW (1999) The effects of 

fishing on fish habitat. In: Benaka L 

(ed).Fish habitat: essential fish habitat 

and rehabilitation. American Fisheries 

Society, Symposium 22, Bethesda, 

Maryland. pp. 150-187

Ball B, Munday B, Tuck I (2000) Effects of otter 

trawling on the benthos and environment 

in muddy sediments. In: Kaiser MJ, 

de Groot SJ (eds). Effects of fishing 

on non-target species and habitats: 

biological, conservation and socioeconomic issues. Blackwell Science 

Ltd., Oxford, UK. pp 69-82
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

100

Barnhart JP (2003) Weathervane scallop 

fishery in Alaska with a focus on the 

Westward Region, 1967-2002. Alaska 

Department of Fish and Game, Division 

of Commercial Fisheries, Regional 

Information Report No. 4K-03-5, 

Kodiak, Alaska, 33 pp

Bayer FM (1952) Two new species of Arthrogorgia

(Goronacea [sic]: Primnoidae) from the 

Aleutian Islands region. Proceedings 

of the Biological Society of Washington 

65:63-70

Bayer FM (1982) Some new and old species of 

the Primnoid genus Callogorgia Gray, 

with a revalidation of the related genus 

Fanellia Gray (Coelenterata: Anthozoa). 

Proceedings of the Biological Society of 

Washington 95:116-160

Bayer FM (1996) The gorgonian genus 

Arthrogorgia (Octocorallia: Primnoidae). 

Proceedings of the Biological Society of 

Washington 109:605-628.

Bayer FM, Macintyre IG (2001) The mineral 

composition of the axis and holdfast 

of some gorgonacean octocorals 

(Coelenterata: Anthozoa), with special 

reference to the family Gorgoniidae. 

Proceedings of the Biological Society 

of Washington 114:309–45

Brodeur RD (2001) Habitat-specific distribution 

of Pacific ocean perch (Sebastes 

alutus) in Pribilof Canyon, Bering Sea. 

Continental Shelf Research 21:207-24

Brooke S, Stone R (in press) Reproduction 

of deepwater hydrocorals (Order 

Anthoathecatae) from the Aleutian 

Islands. Bulletin of Marine Science

Cairns SD (1991) The marine fauna of New 

Zealand: Stylasteridae (Cnidaria: 

Hydroida). New Zealand Oceanographic 

Institute Memoir 98:7-95

Cairns SD (1992) Worldwide distribution of the 

Stylasteridae. Scientia Marina 56:125-

130

Cairns SD (1994) Scleractinia of the Temperate 

North Pacific. Smithsonian Contributions 

to Zoology No. 557, 150 pp

Caldeira K, Wickett ME (2003) Anthropogenic 

carbon and ocean pH. Nature 425:365

Churchill JH (1989) The effect of commercial 

trawling on sediment resuspension and 

transport over the Middle Atlantic Bight 

continental shelf. Continental Shelf 

Research 9:841-865

Cimberg RL, Gerrodette T, Muzik K (1981) Habitat 

requirements and expected distribution 

of Alaska coral. Final Report, Research 

Unit 601, VTN Oregon, Inc. Report 

prepared for the Office of Marine 

Pollution Assessment, Alaska Office, 

U.S. Department of Commerce, NOAA, 

OCSEAP Final Report 54:207-308 

Conwell CN (1976) Progress and 

prospects in marine mining in Alaska. 

State of Alaska, Department of Natural 

Resources Report, Information Circular 

22

Dall WH (1884) On some Hydrocorallinae from 

Alaska and California. Proceedings of 

the Biological Society of Washington 

2:111-115

Etnoyer P, Morgan LE (2005) Habitat-forming 

deep-sea corals in the Northeast Pacific 

Ocean. Pages 331-343 in Freiwald A, 

Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Feely RA, Sabine CL, Lee K, Berelson W, 

Kleypas J, Fabry VJ, Millero FJ (2004) 

Impact of anthropogenic CO2 on the 

CACO3 system in the oceans. Science 

305:362-366

Fisher WK (1938) Hydrocorals of the North Pacific 

Ocean. Proceedings of the United 

States National Museum 84:493-554

Fritz LW, Greig A, Reuter R (1998) Catch-per-uniteffort, length, and depth distributions, of 

major groundfish and bycatch species 

in the Bering Sea, Aleutian Islands 

and Gulf of Alaska Regions based on 

groundfish fishery observer data. U.S. 

Department of Commerce, NOAA 

Technical Memorandum NMFS-AFSC 

88

Gislason H (1994) Ecosystem effects of fishing 

activities in the North Sea. Marine 

Pollution Bulletin 29:520-527

Goñi R (1998) Ecosystem effects of marine 

fisheries: an overview. Ocean and 

Coastal Management 40:37-64
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

101

Greene HG, Yoklavich MM, Starr RM, O’Connell 

VM, Wakefield WW, Sullivan DE, McRea 

JE, Cailliet GM (1999) A classification 

scheme for deep seafloor habitats. 

Oceanologica Acta 22:663-678

Guinotte JM, Buddemeier RW, Kleypas JA (2003) 

Future coral reef habitat marginality: 

temporal and spatial effects of climate 

change in the Pacific basin. Coral Reefs 

22:551-558

Guinotte JM, Orr J, Cairns S, Freiwald A, Morgan 

L, George R (2006) Will human-induced 

changes in seawater chemistry alter the 

distributions of deep-sea scleractinian 

corals? Frontiers in Ecology and the 

Environment 4:141-146.

Heifetz J (2002) Coral in Alaska: distribution, 

abundance, and species associations. 

Hydrobiologia 471:19-28

Heifetz J, Wing BL, Stone RP, Malecha PW, 

Courtney DL (2005) Corals of the Aleutian 

Islands. Fisheries Oceanography 14 

(Suppl. 1):131-138

High WL (1998) Observations of a scientist/

diver on fishing technology and 

fisheries biology. U.S. Department of 

Commerce, National Marine Fisheries 

Service, Alaska Fisheries Science 

Center Processed Report 98-01, Seattle 

Washington

Hoff GR, Stevens B (2005) Faunal assemblage 

structure on the Patton Seamount 

(Gulf of Alaska, USA). Alaska Fishery 

Research Bulletin 11(1):27-36

Horowitz WL (2002) Evaluation of sub-sea 

physical environmental data for the 

Beaufort Sea OCS and incorporation 

into a Geographic Information System 

(GIS) database. U.S. Department of the 

Interior, Minerals Management Service, 

Alaska Outer Continental Shelf Region. 

OCS Study MMS 2002-017 

Hughes SE (1981) Initial U.S. exploration of nine 

Gulf of Alaska seamounts and their 

associated fish and shellfish resources. 

Marine Fisheries Review 42:26-33.

Johnson T (2003) The Bering Sea and Aleutian 

Islands: region of wonders. Alaska Sea 

Grant College Program, University of 

Alaska Fairbanks, Fairbanks, Alaska, 

USA 

Jones JB (1992) Environmental impact of trawling 

on the seabed: a review. New Zealand 

Journal of Marine and Freshwater 

Research 26:59-67

Keller NB (1976) The deep-sea madreporarian 

corals of the genus Fungiacyathus

from the Kurile-Kamchatka, Aleutian 

Trenches and other regions of the world 

oceans. Trudy Instituta Okeanologii 

99:31-44

Kinder TH, Schumacher JD (1981) Circulation 

over the continental shelf of the 

southeastern Bering Sea. In: Hood 

DW, Calder JA (eds) The Eastern 

Bering Sea Shelf: oceanography and 

resources, volume I. U.S. Department 

of Commerce, Washington, D.C, USA 

Kleypas JA, Buddemeier RW, Archer D, Gattuso 

J-P, Langdon C, Opdyke BN (1999) 

Geochemical consequences of 

increased atmospheric carbon dioxide 

on coral reefs. Science 284:118-120

Kleypas JA, Feely RA, Fabry VJ, Langdon C, 

Sabine CL, Robbins LL (2006) Impacts 

of ocean acidification on coral reefs 

and other marine calcifiers: a guide for 

future research. Report of a workshop 

18-20 April 2005, St. Petersburg, FL., 

sponsored by NSF, NOAA, and the U.S. 

Geological Survey. 88 pp

Krieger KJ (2001) Coral (Primnoa) impacted by 

fishing gear in the Gulf of Alaska. In: 

Willison et al. (eds) Proceedings of the 

First International Symposium on DeepSea Corals, Ecology Action Centre and 

Nova Scotia Museum, Halifax, Nova 

Scotia, pp 106-116

Krieger KJ, Wing BL (2002) Megafaunal 

associations with deepwater corals 

(Primnoa spp.) in the Gulf of Alaska. 

Hydrobiologia 471:83-90 

Lamb A, Hanby BP (2005) Marine Life of the Pacific 

Northwest: a photographic encyclopedia 

of invertebrates, seaweeds and selected 

fishes. Harbour Publishing, Madeira 

Park, British Columbia. 398 pp
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

102

Langdon C, Broecker WS, Hammond DE, Glenn 

E, Fitzsimmons K, Nelson SJ, Peng 

T-H, Hajdas I, Bonani G (2003) Effect 

of elevated CO2 on the community 

metabolism of an experimental coral 

reef. Global Biogeochemical Cycles 

17(1):1-14 

Langdon C, Atkinson MJ (2005) Effect of 

elevated pCO2 on photosynthesis and 

calcification of corals and interactions 

with seasonal change in temperature/

irradiance and nutrient enrichment. 

Journal of Geophysical Research 110:

C09S07, doi:10.1029/2004JC002576

Levings CD, McDaniel NG (1974) A unique 

collection of baseline biological data: 

benthic invertebrates from an underwater cable across the Strait of Georgia. 

Fisheries Research Board of Canada 

Technical report No. 441. 19 pp

Lindholm JB, Auster PJ, Kaufman LS (1999) 

Habitat-mediated survivorship of 

juvenile (0-year) Atlantic cod Gadus 

morhua. Marine Ecology Progress 

Series 180:247-255 

Masuda MM, Stone RP (2003) Biological 

and spatial characteristics of the 

weathervane scallop Patinopecten 

caurinus at Chiniak Gully in the central 

Gulf of Alaska. Alaska Fishery Research 

Bulletin 10:104-118

Malecha PW, Stone RP, Heifetz J (2005) Living 

substrate in Alaska: distribution, 

abundance and species associations. 

In: Barnes PB, Thomas JP (eds) 

Benthic habitats and the effects of 

fishing. American Fisheries Society, 

Symposium 41:289-299

Maloney NE (2004) Sablefish, Anoplopoma 

fimbria, populations on Gulf of Alaska 

seamounts. Marine Fisheries Review 

66(3):1-12 

National Academy of Sciences (1990) Assessment 

of the U.S. Outer Continental Shelf 

Environmental studies: 1. Physical 

Oceanography. National Academy 

Press, Washington, D.C., USA

NMFS (National Marine Fisheries Service) (2004) 

Final programmatic supplemental 

groundfish environmental impact 

statement for Alaska groundfish 

fisheries. U.S. Department of 

Commerce, NOAA, NMFS; Alaska 

Region, P.O. 21668, Juneau, Alaska 

99802-1668

NMFS (National Marine Fisheries Service) (2005) 

Final environmental impact statement 

for essential fish habitat identification 

and conservation in Alaska. U.S. 

Department of Commerce, NOAA, 

NMFS; Alaska Region, P.O. 21668, 

Juneau, Alaska 99802-1668

Normark WR, Carlson PR (2003) Giant 

submarine canyons: Is size any clue to 

their importance in the rock record? In: 

“Chan MA, Archer AW (eds) Extreme 

depositional environments: mega end 

members in geologic time. Geological 

Society of America, Special Paper 370: 

175-196

NRC (National Research Council) (2002) Effects 

of trawling and dredging on seafloor 

habitat. National Academy Press. 

Washington D.C. 126 pp

NURP (National Undersea Research Program) 

(1996) Gulf of Alaska Initiative. 

Report of West Coast National 

Undersea Research Center Workshop. 

Anchorage, Alaska, USA

NURP (National Undersea Research Program) 

(2001) ALVIN/Gulf of Alaska Workshop. 

University of Washington, Seattle, 

Washington, USA

Nutting CC (1912) Descriptions of the Alcyonaria 

collected by the U.S. Fisheries steamer 

“Albatross,” mainly in Japanese waters, 

during 1906. Proceedings of the United 

States National Museum 43:1-104, 

plates 1-21 

O’Connell VC, Brylinski C, Greene HG (in press) 

The use of geophysical survey data in 

fisheries management: a case history 

from Southeast Alaska. In: Todd B, 

Greene HG (eds) Marine Benthic 

Mapping. Geological Association of 

Canada Special Paper 47
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

103

Opresko DM (2005) New genera and species 

of antipatharian corals (Cnidaria: 

Anthozoa) from the North Pacific.

Zoologische Mededelingen, Leiden 79-

2 (7):129-165

Orr JC, Fabry VJ, Aumont O, Bopp L, Doney SC, 

Feely RA, Gnanadesikan A, Gruber N, 

Ishida A, Joos F, Key RM, Lindsay K, 

Maier-Reimer E, Matear R, Monfray 

P, Mouchet A, Najjar RG, Plattner GK, Rodgers KB, Sabine CL, Sarmiento 

JL, Schlitzer R, Slater RD, Totterdell IJ, 

Weirig M-F, Yamanaka Y, Yool A (2005) 

Anthropogenic ocean acidification 

over the twenty-first century and its 

impact on calcifying organisms. Nature 

437:681-686

Roberts JM, Wheeler AJ, Freiwald A (2006) Reefs 

of the deep: the biology and geology of 

cold-water coral ecosystems. Science 

312:543-547

Robertson T (2000) Overview of pipeline 

regulatory requirements in Cook 

Inlet, Alaska. Report prepared for the 

Cook Inlet Regional Citizens’ Advisory 

Council, Anchorage, Alaska, 31 pp

Royer TC (1981) Baroclinic transport in the Gulf 

of Alaska, Part I. Seasonal variations of 

the Alaska Current. Journal of Marine 

Research 39:239-250

Rose CS, Jorgensen EM (2005) Spatial and 

temporal distributions of trawling off 

Alaska: consideration of overlapping 

effort when evaluating the effects of 

fishing on habitat. In: Barnes PB, 

Thomas JP (eds) Benthic habitats 

and the effects of fishing. American 

Fisheries Society, Symposium 41:679-

690

Seibel BA, Walsh PJ (2001) Potential impacts 

of CO2 injection on deep-sea biota. 

Science 294:319-320

Shirley SM, Kruse GH (1995) Development of 

the fishery for weathervane scallops, 

Patinopecten caurinus (Gould, 1850), 

in Alaska. Journal of Shellfish Research 

14:71-78

Shotwell SK, Heifetz J, Courtney DL, Greene 

HG (in press) Mapping marine benthic 

habitat in the Gulf of Alaska: geological 

habitat, fish assemblages, and fishing 

intensity. In: Todd B, Greene HG (eds) 

Marine Benthic Mapping. Geological 

Association of Canada Special Paper 

47 

Stone RP, Wing BE (2001) Growth and recruitment 

of an Alaskan shallow-water gorgonian. 

In: Willison et al. (eds) Proceedings 

of the First International Symposium 

on Deep-Sea Corals, Ecology Action 

Centre and Nova Scotia Museum, 

Halifax, Nova Scotia, pp 88-94

Stone RP, Masuda MM, Malecha PW (2005) 

Effects of bottom trawling on softsediment epibenthic communities in the 

Gulf of Alaska. In: Barnes PB, Thomas 

JP (eds) Benthic habitats and the 

effects of fishing. American Fisheries 

Society, Symposium 41:461-475

Stone RP (2006) Coral habitat in the Aleutian 

Islands of Alaska: depth distribution, 

fine-scale species associations, and 

fisheries interactions. Coral Reefs 

25:229-238

Stone RP, Mondragon J, Andrews A (in 

preparation) Deep-sea emergence of 

red tree coral (Primnoa pacifica) in a 

Southeast Alaska glacial fiord

Tomascik T, Mah AJ, Nontji A, Moosa MK (1997) 

The Ecology of the Indonesian Seas. 

Part Two. Periplus Editions, Singapore. 

745 pp 

Turk T (2001) Spatial distribution and selected 

habitat preferences of weathervane 

scallops (Patinopecten caurinus) in 

Alaska. In: Kruse BH, Bez N, Booth A, 

Dorn MW, Hills S, Lipcius RN, Pelletier 

D, Roy C, Smith SJ, and Witherell D 

(eds). Alaska Sea Grant Publication 

No. AK-SG-01-02, University of Alaska, 

Anchorage, pp. 463-477 

Vallier TL, Scholl DW, Fisher MA, Bruns TR, 

Wilson FH, von Huene R, Stevenson 

AJ (1994) Geologic framework of the 

Aleutian arc, Alaska. In: Plafker G and 

Berg HC (eds) The Geology of Alaska. 

The Geology Society of America, Inc., 

Boulder, Colorado, pp 367-388
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

104

Verrill AE (1865) Synopsis of the polyps and 

corals of the North Pacific Exploring 

Expedition, under Commodore C. 

Ringgold and Captain John Rodgers, 

U.S.N., from 1853 to 1856. Collected 

by Dr. Wm. Stimpson, naturalist to the 

Expedition. With descriptions of some 

additional species from the west coast 

of North America. Proceedings of the 

Essex Institute, Salem 4:181-196

Wahle CM (1983) The roles of age, size and 

injury in sexual reproduction among 

Jamaican corals. American Zoologist 

23:961

Whitney FA, Crawford WR, Harrison PJ (2005) 

Physical processes that enhance 

nutrient transport and primary 

productivity in the coastal and open 

ocean of the subarctic NE Pacific. 

Deep-Sea Research II 52:681-706

Wilson MT, Andrews AH, Brown AL, Cordes 

EE (2002) Axial rod growth and age 

estimation of the sea pen, Halipteris 

willemoesi Kölliker. Hydrobiologia 

471:133-142

Wing BL, Barnard DR (2004) A field guide to 

Alaskan corals. NOAA Technical 

Memorandum NMFS-AFSC-146, 67 

pp

Witherell D, Coon C (2001) Protecting gorgonian 

corals off Alaska from fishing impacts. 

In: Willison et al. (eds) Proceedings 

of the First International Symposium 

on Deep-Sea Corals, Ecology Action 

Centre and Nova Scotia Museum, 

Halifax, Nova Scotia, pp 117-125

Witherell D, Woodby D (2005) Application of 

marine protected areas for sustainable 

fisheries production and marine 

biodiversity off Alaska. Marine Fisheries 

Review 67(1):1-27
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Appendix 2.1. Geographical distribution of corals in Alaska waters. EG = eastern Gulf of Alaska, 

WG = western Gulf of Alaska, AI = Aleutian Islands, BS = Bering Sea, SM = seamounts. ● = 

documented, ○ = not documented. Adapted from Heifetz et al. (2005).

Taxa EG WG AI BS SM

Phylum Cnidaria, Class Anthozoa, 

Order Scleractinia (Stony Corals) ● ● ● ● ●

Balanophyllia elegans ● ● ● ○ ○

Caryophyllia alaskensis ● ● ● ○ ○

Caryophyllia arnoldi ● ● ● ○ ●

Crispatotrochus foxi ○ ○ ● ○ ○

Flabellum sp. ○ ○ ● ○ ○

Fungiacyathus marenzelleri ○ ○ ● ○ ○

Fungiacyathus sp. A ○ ○ ○ ○ ●

Javania borealis ○ ● ● ○ ●

Javania cailleti ● ● ● ○ ○

Leptopenus discus ○ ○ ● ○ ○

Paracyathus sp. ○ ○ ● ○ ○

Order Antipatharia (Black Corals) ● ● ● ● ●

Bathypathes alternata ● ○ ○ ○ ○

Bathypathes patula ● ○ ○ ○ ●

Bathypathes sp. A ● ○ ○ ○ ●

Bathypathes sp. B ○ ○ ○ ○ ●

Chrysopathes formosa ● ○ ○ ○ ○

Chrysopathes speciosa ● ○ ○ ○ ○

Dendrobathypathes boutillieri ● ○ ● ○ ●

Heliopathes pacifica ○ ○ ○ ○ ●

Lillipathes lilliei ● ○ ○ ○ ○

Lillipathes wingi ● ○ ○ ○ ●

Parantipathes sp. ● ○ ● ○ ●

Stichopathes sp. ○ ○ ○ ○ ●

Trissopathes pseudotristicha ○ ○ ● ○ ●

Trissopathes tetracrada ○ ○ ○ ○ ●

Subclass Octocorallia, Order 

Alcyonacea, Suborder Alcyoniia (True 

Soft Corals) ● ● ● ● ●

Alcyonium sp. ○ ○ ● ○ ○

Anthomastus japonicus ○ ○ ● ● ○

Anthomastus cf. japonicus ○ ○ ● ○ ○

Anthomastus ritteri ● ● ● ○ ○

Anthomastus sp. A ○ ○ ● ○ ●

Anthothela cf. grandiflora ○ ○ ● ○ ○

Eunephthea rubiformis ● ● ● ● ○

Eunephthea sp. A ○ ○ ● ○ ○

cf. Eunephthea ○ ○ ● ○ ○
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Taxa EG WG AI BS SM

Suborder Stolonifera (Stoloniferans) ● ● ● ● ●

Clavularia armata ○ ○ ○ ○ ●

Clavularia moresbii ● ● ● ○ ○

Clavularia rigida ○ ○ ○ ○ ●

Clavularia sp. A ○ ○ ● ○ ○

Sarcodictyon incrustans ● ○ ● ● ○

Sarcodictyon sp. A ○ ○ ○ ○ ●

Order Gorgonacea (Gorgonians) ● ● ● ● ●

Acanella sp. ○ ○ ○ ○ ●

cf. Acanthogorgia ○ ○ ● ○ ○

Alaskagorgia aleutiana ○ ○ ● ○ ○

Amphilaphis sp. A ○ ○ ● ○ ○

Amphilaphis sp. B ○ ○ ● ○ ○

Amphilaphis sp. C ○ ○ ● ○ ○

Arthrogorgia ijimai ○ ○ ○ ○ ●

Arthrogorgia kinoshitai ● ● ● ● ○

Arthrogorgia otsukai ○ ○ ● ● ○

Arthrogorgia utinomii ○ ○ ● ○ ○

Calcigorgia beringi ● ○ ● ○ ○

Calcigorgia spiculifera ● ● ● ○ ○

Calcigorgia sp. A ○ ○ ● ○ ○

Calyptrophora japonica ○ ○ ○ ○ ●

cf. Chrysogorgia ○ ○ ● ○ ●

Chrysogorgia sp. A ○ ○ ○ ○ ●

Corallium sp. ○ ○ ○ ○ ●

Cryogorgia koolsae ○ ○ ● ○ ○

Fanellia compressa ○ ○ ● ● ○

Fanellia fraseri ○ ● ● ○ ○

Isidella elongata ○ ○ ○ ○ ●

Isidella paucispinosa ● ● ● ○ ○

Isidella sp. A ○ ○ ● ○ ●

Keratoisis profunda ● ● ● ○ ○

Keratoisis sp. A ○ ○ ○ ○ ●

Keratoisis sp. B ○ ○ ○ ○ ●

Lepidisis sp. A ● ● ○ ○ ○

Lepidisis sp. B ○ ○ ○ ○ ●

Muriceides cylindrica ○ ○ ● ○ ○

Muriceides cf. cylindrica ○ ○ ● ○ ○

Muriceides nigra ○ ○ ● ○ ○

Muriceides sp. A ○ ○ ● ○ ○

Narella sp. A ○ ○ ● ○ ○

Narella sp. B ○ ○ ○ ○ ●

Narella sp. C ○ ○ ○ ○ ●

Narella sp. D ○ ○ ○ ○ ●

Narella sp. E ○ ○ ○ ○ ●
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

107

Taxa EG WG AI BS SM

Narella sp. F ○ ○ ○ ○ ●

Paragorgia arborea ● ● ● ● ○

Paragorgia pacifica ● ○ ○ ○ ○

Paragorgia sp. A ○ ○ ○ ○ ●

Paramuricea sp. ○ ○ ● ○ ○

Parastenella sp. A ○ ○ ● ○ ●

Parastenella sp. B ○ ○ ○ ○ ●

Plumarella flabellata ○ ○ ● ○ ○

Plumarella longispina ● ● ● ● ○

Plumarella spicata ○ ○ ● ○ ○

Plumarella spinosa ○ ○ ● ○ ○

Plumarella sp. A ○ ○ ● ○ ○

Primnoa pacifica ● ● ● ● ○

Primnoa pacifica var. willeyi ● ● ● ○ ●

Primnoa wingi ○ ○ ● ○ ○

“Pseudisidella” sp. ○ ○ ● ○ ●

Radicipes verrilli ○ ○ ● ○ ●

Swiftia beringi ○ ○ ● ○ ○

“cf. Swiftia” marki ● ● ● ○ ○

Swiftia pacifica ● ● ● ○ ●

Swiftia simplex ○ ○ ● ○ ●

“cf. Swiftia” sp. A ○ ● ● ○ ○

Thouarella hilgendorfi ○ ○ ● ○ ○

Thouarella striata ○ ○ ● ○ ○

Thouarella superba ○ ○ ● ○ ○

Thouarella sp. A ○ ○ ● ○ ○

Order Pennatulacea (Sea Pens) ● ● ● ● ●

Anthoptilum grandiflorum ○ ○ ● ● ○

Anthoptilum murrayi ○ ○ ● ● ○

Cavernularia vansyoci ○ ○ ● ○ ○

Halipteris californica ● ○ ○ ○ ○

Halipteris willemoesi ● ● ● ● ○

Pennatula phosphorea ● ○ ○ ○ ○

Protoptilum sp. ● ● ○ ○ ○

Ptilosarcus gurneyi ● ● ● ○ ○

Umbellula lindahli ● ● ● ● ○

Virgularia sp. ● ○ ○ ○ ○

Class Hydrozoa, Order Anthoathecatae 

(Stylasterids) ● ● ● ● ○

Crypthelia trophostega ○ ○ ● ● ○

Cyclohelia lamellata ○ ○ ● ● ○

Cyclohelia sp. A ○ ○ ● ○ ○

Distichopora borealis ● ○ ● ○ ○

Distichopora sp. A ○ ○ ● ○ ○

Errinopora nanneca ○ ○ ● ○ ○
STATE OF DEEP CORAL ECOSYSTEMS IN THE ALASKA REGION

ALASKA

108

Taxa EG WG AI BS SM

Errinopora poutalesii ● ● ● ○ ○

Errinopora stylifera ○ ○ ● ○ ○

Errinopora zarhyncha ○ ○ ● ○ ○

Errinopora sp. A ○ ○ ● ○ ○

cf. Stenohelia ○ ○ ● ○ ○

Stylantheca papillosa ○ ○ ● ○ ○

Stylantheca porphyra ● ○ ○ ○ ○

Stylantheca petrograpta ● ○ ● ○ ○

Stylaster alaskanus ○ ○ ● ○ ○

Stylaster brochi ○ ○ ● ○ ○

Stylaster campylecus campylecus ● ● ● ○ ○

Stylaster campylecus parageus ● ○ ○ ○ ○

Stylaster campylecus trachystomus ○ ○ ● ○ ○

Stylaster campylecus tylotus ○ ○ ● ○ ○

Stylaster cancellatus ● ○ ● ○ ○

Stylaster elassotomus ○ ○ ● ○ ○

Stylaster moseleyanus ○ ○ ● ○ ○

Stylaster polyorchis ○ ○ ● ○ ○

Stylaster stejnegeri ○ ○ ● ● ○

Stylaster venustus ● ○ ○ ○ ○

Stylaster verrillii ● ○ ● ○ ○

Stylaster sp. A ○ ○ ● ○ ○
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

109PACIFIC COAST

STATE OF DEEP CORAL ECOSYSTEMS OF THE U.S. PACIFIC COAST:

CALIFORNIA TO WASHINGTON

Curt E. Whitmire and M. Elizabeth Clarke

I. INTRODUCTION

The U.S. Pacific coast marine region 

encompasses the continental margin off the 

states of California, Oregon and Washington. 

Deep corals were first reported here in the 1860s 

with descriptions by A.E. Verrill, including two 

stony corals, Balanophyllia elegans (1864) and 

Paracyathus stearnsii (1869), and one stylasterid, 

Allopora californica (1866). Because all three 

of the species occur in shallow waters, it is not 

surprising that they were the first in the region 

to be reported (Cairns 1994; Ostarello, 1973). 

Dall (1884) also provided early descriptions of 

hydrocorals off California and Alaska as well as 

accounts from fishermen of bycatch of Stylaster

sp. off the Farallone Islands, California as early 

as 1873.

In addition to the taxonomic literature, records 

of deep corals in the region come from a variety 

of other sources including catch records from 

regional bottom trawl surveys, bycatch data 

collected by fishery observers and observations 

from underwater vehicles (e.g., submersibles 

and remotely operated vehicles (ROVs)). In the 

early 1970s, the Alaska Fisheries Science Center 

(AFSC) began conducting triennial bottom trawl 

surveys of demersal fishes in the region. Early 

surveys included records of pennatulaceans and 

a few gorgonians. Unfortunately, identifications 

down to any appreciable taxonomic level were 

initially very limited, typically only to order or 

sometimes family. In 1998, the Northwest 

Fisheries Science Center (NWFSC) began 

conducting annual bottom trawl surveys, but like 

the early AFSC surveys, identifications of corals 

were initially not a priority. 

Beginning in the 1960s and continuing into the 

current period, there has been an increasing use 

of drop camera systems, submersibles and, most 

recently, ROVs to make in situ observations. 

These underwater photographic platforms have 

been used to explore numerous seafloor features 

in the region including rocky banks (e.g., Pearcy 

et al 1989; Stein et al. 1992; Pirtle 2005; Tissot et 

al. 2006), canyons (e.g., Yoklavich et al. 2000), 

escarpments (e.g., Carey et al. 1990; Clague et 

al. 2001), seamounts (e.g., DeVogelaere 2005) 

and other rocky features (Hyland et al. 2004; 

Brancato et al. 2007). Many of these features 

are known to support deep coral communities. 

The disparate data sources that will be referenced 

throughout this report have their strengths 

and weaknesses as they inform discussions 

of deep coral communities and their habitats. 

For example, trawl surveys in the region are 

limited to low relief, sedimentary habitats that 

support relatively few emergent epifauna as 

compared with hard-bottom habitats (Figure 

3.1). Consequently, bottom trawls rarely sample 

stony corals and stylasterids, but have resulted 

in numerous records of pennatulaceans as well 

as fewer records of gorgonians, black corals 

and soft corals (Table 3.1). Furthermore, the 

level of identification during trawl surveys varied 

according to the priority given for sampling of 

invertebrates and the level of expertise of the 

biologists onboard. Despite these limitations, 

trawl surveys encompassed large portions of 

the continental margin (including shelf and 

slope depths). Consequently, they contribute 

to discussions of general zoogeography of 

some higher coral taxa (e.g., order and family 

levels). On the other hand, in situ photographic 

surveys in the region primarily target rocky, high 

relief structures that support diverse benthic 

communities, many of which include corals. 

Surveys of this type, while limited in extent, 

provide detailed information about the size, 

health and habitat affinities of corals, and their 

relationships between other invertebrates and 

Fishery Resource Analysis & Monitoring Division

Northwest Fisheries Science Center

National Marine Fisheries Service

2725 Montlake Blvd. East 

Seattle, WA 98112-2097
PACIFIC COAST

110

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Table 3.1. General statistics on deep corals sampled during National Marine Fisheries Service (NMFS) bottom 

trawl surveys, which were conducted off the coasts of Washington, Oregon and California by the Alaska and 

Northwest Fisheries Science Centers between 1980 and 2005. A total of 10,526 trawl catch records were 

queried.

# Trawls with Corals % Trawls with Corals % Coral Records

Pennatulaceans 1683 16.0% 74.5%

Gorgonians 202 1.9% 8.9%

Antipatharians 197 1.9% 8.7%

Alcyonaceans 150 1.4% 6.6%

Scleractinians 26 0.2% 1.2%

Stylasterids 1 <0.1% <0.1%

Total 2259 100.0%

demersal fishes. One limitation of photographic 

surveys is the challenge of making specieslevel identifications, however some platforms 

(e.g., submersibles, ROVs) provide specimen 

collection capabilities.

Records of deep corals off the U.S. Pacific coast 

span latitudes between 32 and 49°N and from 

the shoreline out to the seaward boundary of the 

exclusive economic zone (EEZ), including depth 

zones from the intertidal down to the bathyal 

(3900 meters). In total, 101 species of corals 

from six cnidarian orders have been identified 

within the EEZ including 18 species of stony 

corals (Class Anthozoa, Order Scleractinia) from 

seven families, seven species of black corals 

(Order Antipatharia) from three families, 36 

species of gorgonians (Order Gorgonacea) from 

10 families, eight species of true soft corals (Order 

Alcyonacea) from three families, 27 species 

of pennatulaceans (Order Pennatulacea) from 

eleven families, and five species of stylasterid 

corals (Class Hydrozoa, Order Anthoathecatae, 

Family Stylasteridae; Appendix 3.1). 

This chapter includes discussion of deep 

corals in the region and their communities. 

Brief descriptions of regional geology and 

oceanography set the stage for discussions 

of coral zoogeography and general habitat 

characteristics. The authors attempted to identify 

all taxa known to occur in the region, and taxa 

that provide vertical structure as potential habitat 

for other organisms are highlighted. In addition, 

the chapter includes a review of potential impacts 

to corals in the region and related conservation 

measures enacted to protect coral communities 

Figure 3.1. Distribution of general seafloor lithologies transected by bottom trawls conducted by the 

Northwest Fisheries Science Center (NWFSC) from 

2001-2003 during surveys of groundfish off the coast 

of Washington, Oregon and California. Only trawls 

where the position of the fishing gear was known 

were used for analysis. Lithology information for 

the continental margin off Washington and Oregon 

was provided by the Active Tectonics and Seafloor 

Mapping Lab at Oregon State University. Lithology 

information for off California was provided by the 

Center for Habitat Studies at Moss Landing Marine 

Laboratories.

R elative F req u en cy

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%

Roc k y S edim entary
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

111PACIFIC COAST

and their habitats. Finally, the authors provide 

recommendations for future research to improve 

our understanding of these organisms.

II. GEOLOGICAL SETTING 

The marine region off the coasts of Washington, 

Oregon and California accounts for about 7% 

(778,628 km2) of the total area of the U.S. EEZ 

(NMFS 2007 in prep). The continental margin in 

this region is characterized by a narrow (5-40 km) 

shelf and steep continental slope, with the shelf 

break at approximately 200 meters water depth. 

The outer continental shelf off Oregon and parts 

of California are marked by large rocky banks 

(Figure 3.2), some of which were at or near the 

surface during the lower sea level stands of the 

glacial epochs. Several of these banks as well 

as other bathymetric features such as pinnacles 

and seamounts create localized upwelling 

conditions that concentrate nutrients, thus driving 

a high level of biologic productivity. For example, 

Heceta Bank, which rises over 100 meters above 

the edge of the continental shelf and to within 70 

meters of the ocean surface, diverts the main flow 

of the California Current, introducing eddies and 

other instabilities that affect areas downstream 

and along the Oregon coast. Smaller rocky 

banks are located off southern California in 

what is called the continental borderlands, a 

geologically complex region characterized by 

deep basins and elevated ridges, some of which 

breech the surface to form the Channel Islands. 

Throughout the region, many of these high relief, 

bathymetric features have been found to support 

coral communities (see Love et al. 2007; Tissot 

et al. 2006; Tissot et al. in prep; Strom 2006). 

The shelf and slope are cut by numerous 

submarine canyons, including one of the deepest 

and largest on the west coast of North America 

- Monterey Canyon (Figure 3.2). Deep corals 

have been discovered at numerous sites here, 

and also within another large canyon – Astoria 

– located directly off the mouth of the Columbia 

River. Other major canyons include Juan de Fuca, 

Quinault, Bodega, Pioneer and Sur Canyons. 

Beyond the slope, a number of seamounts rise 

above the abyssal plain including Thompson, 

President Jackson, San Juan, Rodriquez, Taney, 

Guide, Pioneer and Davidson (Figure 3.2). The 

seamounts off the west coast of North America 

have formed over the past tens of millions of 

years by hotspot volcanism and by enhanced 

melting in association with the migration of the 

spreading centers over a heterogeneous mantle 

(Davis and Karsten 1986). One of the largest 

seamounts – Davidson – has been the site of 

several ROV surveys that have discovered a 

diverse coral community (see DeVogelaere 

2005). A number of additional seamounts that 

are known to support deep corals lie just to the 

west of the EEZ boundary, including Cobb, Brown 

Bear, Fieberling and Jasper.

Other megascale (i.e., km to 10s of km, Greene 

et al. 1999) structural features in the region 

that contribute hard-bottom habitats include the 

Mendocino and Gorda Ridges (Figure 3.2). The 

Mendocino Fracture Zone is a 3000-km long 

transform fault extending from Cape Mendocino, 

California across the Pacific Plate. A prominent 

hard-bottom feature associated with this fault 

is the Mendocino Ridge, which shoals to 1100 

meters water depth and drops 2100 meters to 

the north-south trending Gorda Ridge. To the 

south, the Mendocino ridge slopes 3300 meters 

to the abyssal plain. In contrast to the Mendocino 

Ridge, the Gorda Ridge is a seafloor spreading 

center where two plates are moving apart, 

allowing molten magma to rise up to form new 

oceanic crust. It extends 300 km and is bounded 

by the Mendocino Fracture Zone to the south 

and the Blanco Fracture Zone to the north. The 

Gorda Ridge rises to a maximum height of >1500 

meters above the axial valley floor, which ranges 

from 3200 to >3800 meters water depth. Like 

other spreading centers, hydrothermal vents that 

support unique communities of chemosynthetic 

organisms exist nearby. Morphology of the 

Mendocino and Gorda Ridges were described by 

Fisk et al. (1993) and Clague and Holmes (1987), 

respectively.

III. OCEANOGRAPHIC SETTING

Oceanographic circulation in the region is well 

described and was summarized recently by 

NMFS (2007 in prep). The U.S. Pacific region 

is one of the major upwelling areas of the world, 

where nutrient-rich waters support high levels of 

biological productivity. Physical oceanography 

varies seasonally and during El Niño and La 

Nina events or periods of interdecadal climate 

regime shifts. Major oceanographic currents 
PACIFIC COAST

112

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Figure 3.2. Bathymetric map of the U.S. Pacific coast region. Major topographic features that are mentioned 

in this chapter are labeled.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

113PACIFIC COAST

in the region include the surfaceflowing California Current, the Inshore 

Countercurrent (Davidson Current) and 

the Southern California Countercurrent, 

and the subsurface-flowing California 

Undercurrent. The California Current 

forms the eastern boundary of a large 

clockwise circulation pattern in the North 

Pacific. The California Current is a yearround flow that transports cold, nutrientrich subarctic water equatorward (Hickey 

1998). It extends from the shelf break 

to about 1000 km offshore with peak 

speeds at the surface during spring 

and summer yet significant flows down 

to 500 meters (Hickey 1998). South 

of Point Conception, the California 

Current splits, with its core continuing 

farther offshore while a smaller portion 

turns shoreward both north and south 

of the Channel Islands (Figure 3.3). 

Near San Diego, part of the core flow 

turns northward to form the Southern 

California Countercurrent, an inshore 

poleward flow off Southern California. 

During some years, the Southern 

California Countercurrent forms a 

counterclockwise circulation pattern 

known as the Southern California Eddy 

(Figure 3.3). During other years, the 

Southern California Countercurrent 

rounds Point Conception and combines 

with the Inshore Countercurrent, a poleward flow 

inshore of the equatorward California Current 

(Hickey 1979, Figure 3.3). Below these surface 

currents lies a narrow, high speed flow known 

as the California Undercurrent, which brings 

warmer, nutrient-poor waters poleward along the 

slope from the eastern equatorial Pacific (Pierce 

et al. 2000). A major feature of the eastern North 

Pacific and the California Current is a layer where 

oxygen concentrations are low within a depth 

zone along the upper continental slope, extending 

to depths greater than 1000 m. This feature lies 

beneath the California Undercurrent and is called 

the oxygen minimum zone (dissolved oxygen 

<0.5 ml l-1 (22 mM kg-1) (Deuser 1975).

Circulation in the region is driven by the intensity 

and duration of prevailing seasonal winds and 

storms. Spring and summer northwesterly 

winds drive an upwelling system that replenishes 

nutrients to the photic zone, which in turn 

stimulate biological productivity (Batchelder et 

al. 2002). A shift in wind direction in the winter 

north of Santa Cruz, CA creates favorable 

downwelling conditions, while upwelling 

conditions persist year-round south of Santa Cruz 

due to modest storm activity (Strub and James 

2000). Upwelling plumes also occur at coastal 

headlands, particularly where the California 

Current and Inshore Countercurrent intersect off 

Pt. Conception.

Another important hydrographic feature in the 

northern part of the region is the Columbia 

River plume, contributing as much as 90% of 

the freshwater input between the Strait of Juan 

de Fuca and San Francisco Bay (NMFS 2007 

in prep). The position of the plume is highly 

seasonal, generally extending equatorward and 

offshore in the spring and summer, and poleward 

along the coast in the fall and winter (Thomas 

and Weatherbee 2006). 

According to Briggs (1974), the temperate 

Figure 3.3. Map of portion of the Pacific coast of North America 

showing major oceanographic currents. Image source: NMFS 

2007, in prep.
PACIFIC COAST

114

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

northeastern Pacific includes three coastal 

zoogeographic provinces, two of which include 

waters off Washington, Oregon and California. 

The San Diego Province, in the warm-temperate 

region, extends from Point Conception, CA south 

to Magdalena Bay, Baja California Sur, Mexico. 

The Oregon Province is the lower boreal province 

in the cold temperate region and extends from 

Point Conception north to the Dixon Entrance, 

the maritime boundary between Alaska and 

British Columbia. Another review of Pacific Ocean 

zoogeography by McGowan (1971) placed the 

northern boundary of the Oregon (or “Oregonian”) 

Province at the Strait of Juan de Fuca. 

IV. STRUCTURE AND HABITAT-FORMING 

DEEP CORALS

Several coral taxa in the region are designated 

as “structure-forming”, meaning they are known 

to provide vertical structure above the sea floor 

that can be utilized by other invertebrates or 

fish. Attributes contributing to structure include 

morphology (i.e., branching vs. non-branching), 

whether or not the taxa is known to form 

reefs, relative abundance, maximum colony 

size, manner of colony spatial dispersion (i.e., 

solitary vs. clumped) and the relative number of 

associations with other organisms (Table 3.2). 

With the exception of maximum colony size, 

structure-forming attributes are specific to records 

of corals in the region. Likewise, discussions of 

structure-forming taxa that follow are limited to 

current information in this region. Taxa that are 

considered structure-forming in other regions may 

not be designated as so in this region due to lack 

of specific information on structural attributes. 

a. Stony corals (Class Anthozoa, Order 

 Scleractinia)

Stony corals off the Pacific coast include 18 

species from seven families (Appendix 3.1). 

Most of the records of stony coral are of solitary, 

non-branching cup corals (e.g., Balanophyllia 

elegans). However, at least seven branching 

species are known to occur in the region, 

including Lophelia pertusa, Oculina profunda, 

Madrepora oculata, Dendrophyllia oldroydae, 

Astrangia haimei, Labyrinthocyathus quaylei

and Coenocyathus bowersi. L. pertusa is widely 

distributed off southern California (Hardin et al. 

1994; Mary Elaine Helix, MMS, pers. comm.) and 

has been collected at four sites in the Olympic 

Coast National Marine Sanctuary (Hyland et 

al. 2004; Brancato et al. 2007). One site in the 

Olympic Coast NMS may support the largest 

aggregation of L. pertusa reported in the northeast 

Pacific. Because of this discovery and the fact 

that L. pertusa is relatively abundant off southern 

California, it was given a high rating of structural 

importance (Table 3.2). Other structure-forming 

stony corals are Dendrophyllia oldroydae and 

Oculina profunda (Stephen Cairns, Smithsonian 

Institution, pers. comm.; Table 3.2).

b. Black corals (Class Anthozoa, Order 

 Antipatharia)

Antipatharians are very abundant off the Pacific 

coast but not very speciose with only seven 

species from three families, Antipathidae, 

Cladopathidae and Schizopathidae (Opresko 

2005, 2003, 2002, Appendix 3.1). Antipathes

sp. and Bathypathes sp. exhibit coast 

wide distributions, while Lillipathes sp. and 

Umbellapathes sp. have only been collected at 

Davidson Seamount (Erica Burton, Monterey 

Bay NMS, pers. comm.). Antipathes sp. and 

Bathypathes sp. are branching, can reach 

heights >30 cm, and occur in high abundance. 

A. dendrochristos (Opresko 2005), a newly 

described species ranging in height from 10 to 

250 cm, has been observed via submersible on 

several deepwater banks off southern California 

at water depths between 90 and 360 meters 

(Love et al. 2007; Tissot et al. 2006; Yoklavich and 

Love 2005). Many of these specimens showed 

epifaunal associations with other invertebrates 

including crinoids, amphipods, brittle stars, 

anemones, sponges and crabs. One large (2.1 m 

high) dead colony, which was heavily colonized by 

over 2500 invertebrate individuals, was recently 

aged to 140 years (Love et al. 2007). For these 

reasons, Antipathes spp. was given a high rating 

of structural importance (Table 3.2).

c. Gold corals (Class Anthozoa, Order 

 Zoanthidea)

Gold corals are very rare in the region, with only 

one unconfirmed record from Davidson Seamount 

(DeVogelaere et al. 2005).

d. Gorgonians (Class Anthozoa, Order 

 Gorgonacea)

Gorgonians are the most speciose group of 

corals off the Pacific coast with 36 species from 

10 families (Appendix 3.1). Paragorgia arborea

is found in high abundance (including extensive 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

115PACIFIC COAST

“forests” observed along several ridges on 

Davidson Seamount [DeVogelaere et al. 2005]), 

can reach heights >1 m and has shown epifaunal 

relationships with numerous other structureforming invertebrates. P. arborea is therefore 

given a high rating of structural importance 

(Table 3.2). Isidella spp. and Keratoisis spp. are 

found coast wide mostly on the continental slope. 

Although both genera can reach heights greater 

than 30 cm, other gorgonians (e.g., P. arborea 

and Primnoa pacifica) in the region can reach 

heights exceeding 1 meter. Therefore, Isidella

and Keratoisis were given a medium rating of 

structural importance (Table 3.2). ROV surveys 

in the Olympic Coast NMS have resulted in 

numerous observations of gorgonians including 

large colonies of P. pacifica, numerous colonies 

of Plumarella longispina and smaller colonies of

Leptogorgia chilensis, Swiftia pacifica, and Swiftia 

beringi at several sites. Colonies of P. pacifica

obtained off La Jolla, CA (north of San Diego) at 

205-234 meters are the southernmost record of 

the species in the Pacific (Cairns and Barnard 

2005). Specimens of Keratoisis and Corallium 

from Davidson Seamount have been aged to 

over 200 and 115 years, respectively (Andrews 

et al. 2005).

e. True soft corals (Class Anthozoa, Order 

 Alcyonacea)

Only eight species of true soft corals from three 

families occur off the Pacific coast (Appendix 

3.1). Anthomastus sp. is abundant and exhibits 

coast wide distributions, while Gersemia sp. has 

been caught primarily on the northern Oregon 

slope during trawl surveys. Alcyonium rudyi and 

Cryptophyton goddarti were described recently 

(1992 and 2000, respectively) off the Oregon 

coast (Cairns et al. 2002). Other than C. goddarti, 

references to Clavularia and Telesto off southern 

California (SCAMIT 2001), and Telestula ambigua

in deep water off central California (Austin 1985), 

Table Key

Attribute Measure

Reef-Building Yes/No

Relative Abundance Low/ Medium/ High

Size (width or height) Small (< 30cm)/ Medium (30cm-1m)/ Large (>1m)

Morphology Branching/ Non-branching

Associations None/ Few (1-2)/ Many (>2)

Spatial Dispersion Solitary/ Clumped

Overall Rating Low/ Medium/ High

Attributes Contributing To Structure

Taxa

ReefBuildingRelative 

Abundance

Maximum 

Colony 

Size Morphology

Associations 

with Other 

Species

Colony 

Spatial 

Dispersion

Overall 

Rating of 

Structural 

Importance

Lophelia pertusa No High Large Branching Many Clumped High

Oculina profunda No Low ? Branching ? ? Low

Dendrophyllia 

oldroydae No Medium Medium Branching ? Clumped Medium

Antipathes 

dendrochristos No High Large Branching Many Solitary High

Bathypathes sp. No High Medium Branching ? Solitary Medium

Isidella sp. No High Medium Branching ? Solitary Medium

Keratoisis sp. No High Medium Branching ? Solitary Medium

Paragorgia arborea No High Large Branching Many Solitary High

Primnoa pacifica No Low Large Branching Many Solitary High

Table 3.2. Structure-forming attributes of some deep coral taxa off the U.S. Pacific coast. Relative abundance 

data was compiled from taxonomic records, in situ photographic surveys, and to a lesser extent bottom trawl 

surveys. Numbers of species associations were quantified from in situ photographic surveys. Size, morphology and other colony attributes were informed by taxonomic descriptions. ‘?’ means insufficient data available 

in the region to comment.
PACIFIC COAST

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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

there are no other data on stoloniferans in the 

region. Because of their small stature, none of 

the true soft corals in the region are considered 

to be structure-forming.

f. Pennatulaceans (Class Anthozoa, Order 

 Pennatulacea)

Pennatulaceans are the most abundant coral 

taxon in the region and have been observed from 

submersibles and ROVs either alone or in groves 

of numerous individuals similar to aggregations 

observed off Alaska (Stone et al. 2005; Brodeur 

2001). They are also the most common coral 

taxon recorded from trawl surveys (Table 3.1). 

To date, 27 species from eleven families are 

known to occur off the Pacific coast (Appendix 

3.1). Stylatula sp., Anthoptilum grandiflorum and 

Umbellula sp. are the most common taxa, all of 

which are found coast wide. Although groves 

of pennatulaceans have been shown to support 

higher densities of some fish species than 

adjacent areas (e.g., Brodeur 2001), they are not 

considered to be structure forming as defined by 

this report.

g. Stylasterid corals (Class Hydrozoa, 

 Order Anthoathecatae, Suborder Filifera)

Lace corals or stylasterid corals off the Pacific 

coast have been observed colonizing moderate 

to high-relief rocky habitats from the intertidal 

down to shelf water depths. Only five species 

from three genera are known to occur in the 

region (Fisher 1938; Cairns 1983; Alberto 

Lindner, pers. comm., Appendix 3.1). Stylaster 

californicus is the only species known from 

the San Diego Province while S. venustus is 

found throughout the Oregon Province. Other 

species that exhibit narrower distributions in the 

Oregon Province include Errinopora pourtalesii, 

Stylantheca porphyra and S. petrograpta (Fisher 

1938; Cairns 1983). The two Stylaster species 

and E. pourtalesii are flabellate (i.e., fan-shaped) 

while Stylantheca is encrusting (Cairns 1983). 

Because most specimens in the region rarely 

exceed 30 cm in height or width, they are not 

considered to be structure-forming. Stylasterid 

corals have rarely been identified in trawl survey 

catches (Table 3.1), most likely because bottom 

trawls do not target the high-relief habitats and 

shallow depths at which stylasterids are typically 

found.

V. SPATIAL DISTRIBUTION OF CORAL 

SPECIES AND HABITATS

Much of the information on general zoogeography 

of corals in the region originates from taxonomic 

records and bottom trawl surveys conducted by 

the National Marine Fisheries Service (NMFS) 

(Appendix 3.1). The Alaska Fisheries Science 

Center (AFSC) conducted regional trawl surveys 

off the Pacific coast from 1971-2001, and the 

Northwest Fisheries Science Center (NWFSC) 

began ongoing surveys in 1998. Identification of 

invertebrates was initially very limited; therefore 

this report focuses on catch records from 1980-

2005. Cumulatively, both surveys covered much 

of the continental shelf and upper slope (10-1600 

m water depth); however, survey effort has been 

spatially and temporally variable. Prior to 2002, 

there was limited trawl survey effort south of Pt. 

Conception (34.5oN). Also, the number of trawls 

conducted during each survey varied from year 

to year. A total of 7252 AFSC and 3274 NWFSC 

trawl catch records were queried for coral 

occurrences. Pennatulaceans were recorded 

in 16% of survey trawls, while all other coral 

taxa occurred in only 5% of trawls (Table 3.1, 

Figure 3.4). In addition to NMFS, the Southern 

California Coastal Water Resource Project 

(SCCWRP) conducted bottom trawls during three 

different continental shelf surveys off southern 

California from 1994 to 2003 (Allen et al. 1998; 

Allen et al. 2002). Catch records from 304 of 957 

(32%) SCCWRP trawls include 316 records of 

pennatulaceans encompassing thirteen species 

from six families, and 225 records of other corals 

encompassing fifteen species from six families. 

In addition to trawl surveys, Etnoyer and Morgan 

(2003) compiled records of observations and 

collections of structure-forming corals off the 

Pacific coast by the California Academy of 

Sciences (CAS), the Monterey Bay Aquarium 

Research Institute (MBARI), the Smithsonian’s 

National Museum of Natural History (NMNH), 

and Scripps Institution of Oceanography (SIO). 

Taxonomists have confirmed the identities of 

some of these records, but many records are 

limited to higher taxa (e.g., genus and family). 

Records off Washington, Oregon and California 

include representatives from two families of stony 

corals (Oculinidae and Caryophyllidae), one 

family of black corals (Antipathidae), four families 

of gorgonians (Corallidae, Isididae, Paragorgiidae 

and Primnoidae) and stylasterid corals (Family 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

117PACIFIC COAST

Figure 3.4. Maps of frequency of occurrence for two groups of deep coral taxa sampled during NMFS bottom 

trawl surveys (1980-2005). Frequency defined as number of trawls with corals identified in the catch sample 

divided by total number of trawls within each 20x20 km cell. Frequency was categorized into three classes: 

>0-20%, >20-50%, and >50%. Cells where survey trawls occurred but where no corals were identified in the 

catch sample are labeled as “NO Catch” and are symbolized with an empty box where the underlying bathymetry shading is visible. Pennatulaceans were singled out because they inhabit different habitat types and were 

caught much more frequently than other coral taxa. See Table 3.1 for frequency information.
PACIFIC COAST

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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Stylasteridae). A total of 389 records of corals 

span much of the EEZ off the U.S. Pacific coast. 

This information contributes to the general 

zoogeography of coral taxa in the region, many 

of which are highlighted in the following sections 

pertaining to the two zoogeographic provinces.

More detailed information on coral habitats in 

the region is provided by in situ photographic 

surveys. When possible, the data sources and 

brief descriptions of these surveys are provided.

San Diego Province

The U.S. portion of the San Diego Province 

extends from the Mexican border, north to Point 

Conception, CA, and includes the geologically Figure 3.5. Map showing locations of black corals (Order Antipatharia) from NWFSC, AFSC, and SCCWRP trawl surveys, Etnoyer and Morgan (2003) and Tissot et al. (2006). Specific identities from trawl survey catch records are unconfirmed and primarily limited to genus or family level.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

119PACIFIC COAST

complex borderlands (Figure 3.2). A number of 

species found in this region, such as the newly 

described black coral, Antipathes dendrochristos

(Opresko 2005) have not been described further 

north. These black corals have been observed via 

submersible on numerous rocky outcrops in the 

province at water depths ranging from 90 to 360 

m (Love et al. 2007; Tissot et al. 2006; Yoklavich 

and Love 2005, Figure 3.5). Lophelia pertusa

and Desmophyllum dianthus have been observed 

on numerous high-relief, hard-bottom features 

below 120 meters near oil platforms surveyed in 

the late 1980s off Pt. Conception, CA (Steinhauer 

and Imamura 1990; Hardin et al. 1994). Near 

one platform, D. dianthus and L. pertusa were 

among the most abundant taxa observed in high- Figure 3.6. Map showing locations of stony corals (Order Scleractinia) from NWFSC, AFSC, and SCCWRP trawl surveys, Etnoyer and Morgan (2003) and Brancato et al. (in review). Specific identities from trawl survey catch records are unconfirmed and primarily limited to genus or family level. 
PACIFIC COAST

120

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

relief habitats at water depths ranging from 160-

212 meters. Another scleractinian, Caryophyllia 

arnoldi, is known to occur throughout the 

province especially around the Channel Islands 

(Cairns 1994). The cup coral, Paracyathus 

stearnsii, is also common around the Channel 

Islands including 25 specimens deposited at SIO 

(Cairns 1994). Coenocyathus bowersi has been 

collected from the nearshore off the coasts of 

mainland California, around the Channel Islands 

and down to 80 meters off Pt. Conception (Cairns 

1994). One record of the colonial scleractinian, 

Madrepora oculata, collected at 84 meters water 

depth near Anacapa Island, is only one of two 

records known from the northeast Pacific (Cairns 

1994, Figure 3.6). 

Figure 3.7. Map showing locations of gorgonians (Order Gorgonacea) from NWFSC, AFSC, and SCCWRP trawl surveys, Etnoyer and Morgan (2003) and Tissot et al. (2006). Specific identities from trawl survey catch records are unconfirmed and primarily limited to genus or family level.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

121PACIFIC COAST

Gorgonians are not as prevalent as they are in 

the Oregon Province (Figure 3.7); however, this 

may be due to sampling bias. Tissot et al. (2006) 

observed 27 specimens from four different 

habitat types at 144-163 m. Other records in 

the province include Lepidisis sp. at 950 m and 

Keratoisis sp. far offshore San Diego at 3180 and 

3880 m (Etnoyer and Morgan 2003). Gorgonian 

catches from trawl surveys range in water depths 

from 77-1400 meters. Pennatulaceans (mostly 

members of suborder Subselliflorae) have 

been observed from underwater vehicles in the 

sedimented flanks of numerous rocky outcrops in 

the province (9726 specimens from Tissot et al. 

2006) and are caught more often than other coral 

taxa in bottom trawls at water depths ranging Figure 3.8. Map showing locations of true soft corals (Order Alcyonacea) from NWFSC, AFSC, and SCCWRP trawl surveys and Et- noyer and Morgan (2003). Specific identities from trawl survey catch records are unconfirmed and primarily limited to genus or family level.
PACIFIC COAST

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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

from 44 to over 1500 meters. The only stylasterid 

coral known to occur in the province is Stylaster 

californicus, which has been recorded in rocky 

habitats down to 90 meters water depth.

Oregon Province

The Oregon Province, which extends from Pt. 

Conception, CA north to the maritime boundary 

between Alaska and British Columbia, and 

includes one of the more recent discoveries of 

structure-forming stony coral off the U.S. Pacific 

coast – Lophelia pertusa at the Olympic Coast 

NMS. During ROV surveys, L. pertusa was 

observed on a rock ledge in 271 meters of water in 

2004 (Hyland et al. 2004) and at three other sites 

in 2006, including a broad (tens of meters wide), 

low-lying mound (<1 meter high) at approximately 

250 meters water depth (Brancato et al. 2007). 

Both dead and living colonies were observed. 

Desmophyllum sp. was also observed in 

association with L. pertusa (Brancato et al. 2007), 

which is consistent with observations of Lophelia

elsewhere in the region and world (see Hardin 

et al. 1994; Cairns and Stanley 1982). Colonies 

of the stylasterid coral, Stylaster venustus, were 

observed in water depths of about 100 meters off 

Cape Flattery, WA (A. Lindner, pers. obs.). 

Scleractinians and stylasterids are also found 

elsewhere in the province. Labyrinthocyathus 

quaylei is found on Cordell Bank and south to San 

Diego at water depths of 37-293 m (Cairns 1994). 

Stylasterids and solitary cup corals are common 

primarily in nearshore hard bottom habitats 

(Cairns 1983, 1994), including over 36,000 

observations of Balanophyllia elegans at Cordell 

Bank (Pirtle 2005). During surveys of proposed 

submarine cable routes off Pt. Arguello, CA, 

epifaunal coverage of Stylantheca porphyra, two 

cup corals (B. elegans and Paracyathus stearnsii) 

and two species of anemones often reached 100 

percent on moderate-relief hard-bottom habitats 

(Aspen 2006). Caryophyllia arnoldi is known 

from throughout the Oregon Province including 

inside waters of Washington state and British 

Columbia. Paracyathus stearnsii is common in 

Monterey Bay including 45 specimens deposited 

at CAS (Cairns 1994). The Farallone Islands 

(off San Francisco, CA) mark the northern and 

southern distributional extents of two stylasterid 

corals, Stylaster californicus and S. venustus, 

respectively. Other stylasterids include Errinopora 

pourtalesii, which is found off central California at 

shelf depths, and Stylantheca petrograpta, which 

is found only in the northern part of the province 

off the southern part of Vancouver Island and in 

Puget Sound (Fisher 1938; Alberto Lindner, pers. 

comm.).

Black corals in the Oregon Province are most 

prevalent north of Cape Mendocino with the 

largest reported catches from the northern Oregon 

slope (Figure 3.5). Chrysopathes speciosa

and Antipathes sp. are the most common 

antipatharian taxa in the province and entire 

region. Gorgonians are common at all depths 

covered by regional trawl surveys but also more 

common north of Cape Mendocino (Figure 3.6). 

They have also been recorded from water depths 

over 2900 m off San Francisco, CA (Etnoyer and 

Morgan 2003). During submersible dives off the 

Oregon coast in the mid-1990s, high densities 

of gorgonians were observed at depths between 

200 and 250 meters, where hard rocky substrate 

was covered with thick, hummocky sediments at 

the southern edges of submerged rocky banks 

(Strom 2006). Despite being the second-most 

abundant invertebrate taxa observed (21% of all 

organisms encountered), gorgonians observed at 

these sites did not exceed 30 cm in height. The 

soft coral, Anthomastus sp., has been caught 

over all latitudes but only on the slope (Figure 

3.8). Pennatulaceans are ubiquitously distributed 

throughout the region both by latitude and depth 

(Figure 3.9). Most specimens are Stylatula sp., 

but Anthoptilum grandiflorum, Ptilosarcus gurneyi

and Umbellula sp. are also abundant and widely 

distributed. As in the San Diego Province, they 

prefer sedimentary habitats and Stylatula has 

been observed in dense groves. 

VI. SPECIES ASSOCIATIONS WITH DEEP 

CORAL COMMUNITIES

Several studies both in the region and elsewhere 

in the north Pacific report fine-scale associations 

between demersal fishes, corals and other 

structure-forming invertebrates (e.g., Stone 2006; 

Krieger and Wing 2002; Brodeur 2001; Hardin et 

al. 1994; Hixon et al. 1991) and some studies 

have even investigated the nature of those 

relationships (e.g., Tissot et al. 2006; Auster 

2005; Pirtle 2005; Parrish 2004; Syms and Jones 

2001). In addition to in situ observations, corals 

have been collected with other invertebrates 

and fishes during bottom trawl surveys and 

commercial fishing operations. Unfortunately, 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

123PACIFIC COAST

bottom trawls are of limited precision because 

they often extend over kilometers of seabed, 

traversing a variety of low-relief habitats. 

Furthermore, trawl gears are not designed to 

target sessile invertebrates. Therefore, catches 

of corals only represent presence data, and 

cannot be used to develop standardized indices of 

coral abundance. Lastly, catches of corals often 

consist of partial colonies or skeletal fragments, 

making it difficult to ascertain the size and overall 

health of the organism. Consequently, the best 

source of data on species associations comes 

from direct observations via submersibles or 

other in situ photographic methods (e.g., ROVs). 

Three studies in particular have examined the 

nature of relationships between deep corals and Figure 3.9. Map showing locations of pennatulaceans (Order Pennatulacea) from NWFSC, AFSC, and SCCWRP trawl surveys. Specific identities from trawl survey catch records are unconfirmed and primarily limited to genus or family level.
PACIFIC COAST

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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

other fauna in the region (Tissot et al. 2006; Pirtle 

2005; Hardin et al. 1994), though other recent 

video surveys (e.g., Brancato et al. 2007; Tissot 

et al. in prep) are currently being analyzed in this 

context.

San Diego Province

Data analysis of submersible observations from 

11 rocky banks off southern California (Tissot 

et al. 2006) includes the most comprehensive 

study of fish-invertebrate relationships to date 

in the province. Densities of gorgonians and 

black corals were highest in low relief, mixed 

rock areas including boulders, cobbles, pebbles 

and sand. Approximately 15% of the 135 black 

corals observed had other organisms lying on 

or attached to them and epifauna were mostly 

crinoids (6.8%) followed by sponges (3.1%). 

Only 1.3% of all black coral associations involved 

fishes. None of the 27 observations of gorgonians 

showed evidence of epifaunal associations. 

Fish species exhibiting higher frequencies of 

occurrence near large corals and sponges than 

elsewhere along transects included cowcod 

(Sebastes levis; median distance = 5.5 m), bank 

rockfish (S. rufus; 1.0 m), swordspine rockfish 

(S. ensifer; 1.3 m), shortbelly rockfish (S. jordani; 

1.5 m), pinkrose rockfish (S. simulator; 1.7 m) 

and other members of the rockfish subgenus 

Sebastomus (1.4 m). Most of these species 

including cowcod, bank rockfish and several 

members of the subgenus, Sebastomus, have 

been targeted commercially. Although these six 

fish species and large invertebrates co-occurred 

in similar habitats, the authors questioned the 

existence of functional relationships because the 

median distances between them were not small 

(1.0-5.5 m). In addition to fishes, black corals 

(Antipathes sp.) were observed in association 

with crinoids, sponges, crabs, basketstars, 

brittlestars, anemones, algae and salps. 

Hardin et al. (1994) examined the spatial 

distributions of epifauna on both low- and highrelief hard bottom features off Pt. Conception. 

The second-strongest positive correlation 

reported between two taxa was that of the cup 

coral, Desmophyllum dianthus and the colonial 

scleractinian, Lophelia pertusa. Moderately 

positive correlations existed between two 

other cup corals – Paracyathus stearnsii and 

Caryophyllia sp. Other associations with D. 

dianthus included galatheid crabs and anemones 

(e.g., Amphianthus californica). The authors 

suggest that these correlations, while statistically 

significant, are most likely due to common 

affinities to the physical attributes of their habitat 

(depth, relief, orientation to currents), and not 

indicative of any functional relationships among 

taxa.

Oregon Province

At Cordell Bank, Pirtle (2005) examined 

relationships between structure-forming 

megafaunal invertebrates and demersal fishes. 

During submersible dives at water depths from 

55 to 250 meters, gorgonians were observed 

in mixed rock habitats of varying relief while 

pennatulaceans (Ptilosarcus sp. and members 

of suborder Subselliflorae) occurred in lowrelief habitats of mud and bedrock. Widow (S. 

entomelas), rosy (S. rosaceus) and unknown 

juvenile rockfish, adult Sebastomus, black-eyed 

gobies and combfishes were observed more 

often near gorgonians than expected by chance in 

habitats where they occurred. In sand- and muddominated habitats, flatfish, poachers, combfishes 

and greenspotted rockfish (S. chlorostictus) were 

observed near pennatulaceans (Ptilosarcus sp.). 

Greater than half of all associations between 

gorgonians and fishes were within a distance 

of 1 m; however, none involved direct physical 

contact. 

Davidson Seamount, in contrast to Cordell 

Bank, is a much deeper submarine environment. 

Davidson rises from the surrounding seafloor 

at 3600 meters water depth to its crest at 1250 

meters. Corals were observed here primarily on 

ridges and often oriented to maximize surface area 

towards the current (DeVogelaere et al. 2005). 

Rattails (Coryphaenoides sp.) and thornyheads 

(Sebastolobus sp.) were observed adjacent to 

corals, along with sponges, other corals, sea stars, 

clams, sea cucumbers and octopi (Graneldone 

sp.). Coral epifauna included polychaete worms, 

isopods, shrimps, crabs, basket stars, crinoids, 

brittle stars and anemones. 

With the exception of Pirtle (2005), DeVogelaere 

et al. (2005) and Tissot et al. (in prep), much 

of the data from underwater surveys in the 

Oregon Province have yet to be analyzed for 

species associations with deep corals. Recent 

in situ studies at Davidson Seamount, Monterey 

Canyon and in the Olympic Coast NMS may 

elucidate additional species associations. For 

example, the recent ROV survey of deep-sea 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

125PACIFIC COAST

coral assemblages at the OCNMS (Brancato et 

al. 2007) revealed many species of fishes and 

large invertebrates (e.g. shrimp, brittle stars, 

crabs) nestled among the coral structures (see 

also. However, the degree to which corals might 

contribute to the feeding, growth and reproduction 

of demersal fishes or provide biogenic structure 

for other megafaunal invertebrates is largely 

unknown for most of the Oregon Province. 

VII. STRESSORS ON DEEP 

CORAL COMMUNITIES

Compared to other regions in the U.S., the 

Pacific coast from California to Oregon has a 

narrow continental shelf, which may result in 

coral communities here being more susceptible 

to coastal activities. Some activities that may 

adversely effect corals include oil and gas 

development, deployment of gas pipelines and 

communication cables, and marine pollution. 

However, fishing operations, particularly 

bottom trawling, pose the most immediate and 

widespread threats to deep coral communities. 

Effects of fishing

The temperate, nutrient-rich waters of the 

California Current support lucrative commercial, 

tribal and recreational fisheries. Fishing operations 

in the region are very diverse. A limited-entry 

trawl fishery operates from the Canadian border 

south to Morro Bay, CA targeting numerous 

demersal species both on the 

shelf and upper slope. Vessel 

sizes range from 35 to 95 feet 

and average 65 feet (NWFSC 

2006). Off California and 

Oregon, an open-access fishery 

targeting nearshore species is 

comprised of vessels from 10 to 

50 feet in length (NWFSC 2007). 

Charter boats also operate 

out of numerous coastal ports 

and target a variety of pelagic 

and demersal species. Gear 

types used in the region include 

bottom trawls, midwater trawls, 

demersal seines, pots, bottomset gillnets, bottom longlines 

and other hook-and-line gear, 

but bottom trawls are the most 

widely used and potentially 

harmful to corals. The degree of impact to corals 

from fishing operations depends on the physical 

attributes of their habitat (e.g., sediment type, 

relief), attributes of the gear (e.g., configuration, 

mode of operation, footprint) and its geographic 

extent of use (Table 3.3). These attributes will be 

detailed in relation to each gear type used in the 

region.

Bottom trawls

Bottom trawls are the most widely used fishing 

gear off the Pacific coast. They are used off 

Oregon and in federal waters off Washington and 

California to target numerous species of demersal 

fishes, shrimp, prawns, sea cucumbers and sea 

urchins. Bottom trawls also have the greatest 

severity ranking of all gear types used in the 

region (Morgan and Chuendpagdee 2003). Gear 

components that contact the seafloor include the 

doors, bridles, footrope (except in shrimp trawls) 

and occasionally the netting (PFMC 2005). Gear 

configurations depend on target species and 

operating depths, with door separation distances 

ranging from 34-50 m for shelf trawls and 50-

200 m for slope trawls (PFMC 2005). Footrope 

lengths commonly range from 15-34 m for shelf 

fisheries. Due to their weight and speed (1.5-2.5 

knots) over the seafloor, all trawl components that 

contact the seafloor have the potential to snare, 

undercut or topple emergent structures, including 

deep corals. The Pacific Council Groundfish 

Essential Fish Habitat (EFH) Environmental 

Impact Statement (EIS) identified sensitivity 

of coral habitats to trawl gear as relatively high 

Table 3.3. Ratings of potential fishing gear impacts to deep corals off the 

U.S. Pacific coast. Each measure of impact is rated as High (H), Moderate 

(M), or Low (L).

Measure of Impact

Gear Type

Severity of Impact

Extent of Impact

Geographic 

Extent of Use in 

Region

Overall Rating of 

Gear Impact

Bottom trawls H H H H

Midwater trawls L L H L

Demersal seines L M L L

Bottom longlines & gillnets M M M M

Pots & Traps M L L L

Other hook and line L L L L
PACIFIC COAST

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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

(2-3) on a scale of 0-3 with three defined as 

“major changes in bottom structure, such as 

re-arranged boulders; large losses of many 

organisms with differences between impact and 

control sites greater than 50% in most measured 

metrics.” Furthermore, bottom trawls accounted 

for over 92% of observed coral bycatch in the 

region between August 2001 and August 2004 

(Figure 3.10). Because bottom trawls are used 

extensively throughout the region and deep coral 

habitats are particularly sensitive to these gears, 

they were given a high rating of impact (Table 

3.3).

Midwater trawls

Midwater trawls are used extensively in the Figure 3.10. Map showing locations of deep coral bycatch recorded by fishery observers in the West Coast Groundfish Observer Program. All observed trips and gear types from August 2001 – August 2004 were queried. Due to limitations of specific identifica- tions, coral bycatch was grouped into two classes: 1) gorgonians and stony corals, and 2) pennatulaceans. Point symbols repre- sent start locations of bottom trawls or longline and pot sets.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

127PACIFIC COAST

Oregon Province to target Pacific hake (Merluccius 

productus). Hake is a schooling, pelagic species 

and therefore vessels using midwater trawls try 

to avoid contact with the seafloor. As evidenced 

by no records of coral bycatch from this gear type 

between August 2001 and August 2004 and their 

very low rating of habitat impact (Morgan and 

Chuendpagdee 2003), midwater trawls have low 

impacts to deep coral communities in the region 

(Table 3.3).

Demersal seines

Demersal seines, also known as Scottish seines, 

are used in nearshore and shelf areas to catch 

flatfish (e.g., sand dabs, Petrale sole, English 

sole) and chillipepper rockfish (PFMC 2005). 

Demersal seines use a large net attached to 

long (hundreds of meters) ropes to herd fish 

on the seafloor. In contrast to trawl gear, their 

lighter weight and slower movement over the 

seafloor cause little disturbance (PFMC 2005). 

Corals most likely to be impacted by this gear 

are pennatulaceans, because demersal seines 

are used in areas of unconsolidated sediments 

where pennatulaceans inhabit. Due to their 

limited use in the region; however, demersal 

seines were given a low rating of impact (Table 

3.3). Furthermore, there are no records of deep 

coral bycatch from this gear type.

Scallop dredges

Although dredges were once used to target 

weathervane scallops, they are now prohibited in 

the region.

Bottom longlines and gillnets 

Bottom-set gillnets are prohibited offshore 

Washington and Oregon and in California state 

waters. In federal waters off California, gillnets 

are used to catch white seabass, bonito, flying 

fish, white croaker, angel shark, California halibut, 

lingcod, mullet and perch (PFMC 2005). Bottom 

longlines, on the other hand, are used throughout 

the region to target sablefish, rockfish, Pacific 

halibut, cabezon, lingcod or dogfish. Bottom 

longlines are composed of weights, hooks and 

a mainline that contact the seafloor. Gillnets 

are anchored by weights and leadlines, which 

weigh about 100 pounds per 100 fathoms of line 

(PFMC 2005). These gears can travel significant 

distances over the seafloor, particularly during 

retrieval when the vessel is not directly over 

the gear, snaring or undercutting emergent 

structures (e.g., corals). According to Morgan 

and Chuendpagdee (2003), bottom longlines 

and gillnets have a medium to low impact on the 

physical and biological components of habitat. 

Furthermore, from August 2001 through August 

2004, gillnets and longlines accounted for less 

than 4% of all observed coral bycatch. However, 

this low percentage may reflect a sampling bias 

to mobile gears or that corals damaged by these 

gears do not reach the surface. Lost longline gear 

has been observed on the seafloor at several 

sites within the Olympic Coast NMS (Hyland et 

al. 2004; Brancato et al. 2007). Nevertheless, 

sensitivity of coral habitats to hook and line gear 

(e.g., longlines) was classified as low to moderate 

(0.3-1.3) on a scale of 0-3 (PFMC 2005). Due to 

this low to moderate sensitivity rating, medium to 

low habitat impact rating, and their moderate use 

in the region, bottom longlines and gillnets were 

given a medium rating of overall impact (Table 

3.3). 

Other fishing gears

Throughout the region, pots (also called traps) 

are used to catch sablefish, Dungeness and 

other species of crab, spot prawns, spiny 

lobster and other finfish. Pots can be single 

(e.g., Dungeness crab) or in a series of up to 50 

attached to a groundline (e.g., sablefish) (PFMC 

2005). The effect of pots on the seabed depends 

on their weight, shape and lateral movement 

during retrieval. If the vessel is not directly above 

the pot being retrieved, significant contact with 

the bottom can occur; however, this movement 

is typically minimized so as not to put excessive 

strain on the line and other equipment (PFMC 

2005). Severity of impact to coral habitats by 

pots and traps was ranked as low to moderate 

(PFMC 2005; Morgan and Chuendpagdee 2003). 

Factoring in their limited use in the region, pots 

were given a low overall rating of impact (Table 

3.3). 

In addition to longlines, other types of hook and 

line gear (e.g., stick gear, rod and reel, jig gears 

and vertical longlines) are used in the region, 

but their impacts to deep corals are most likely 

minimal. Sinkers, hooks and lines can snare 

and damage corals, but the area of seafloor 

contacted is small relative to trawls and dredges. 

Consequently, these hook and line gears were 

given a low rating of impact (Table 3.3).
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Effects of other human activities

In addition to fishing operations, 

sedimentation caused by oil and gas 

development has been shown to be 

detrimental to corals in the region. Other 

human activities that may adversely 

impact corals include the deployment 

of gas pipelines and communication 

cables and pollution. Unfortunately, 

little is known about these potential 

stressors but they are likely to have only 

localized impacts, if any. In the future, 

global climate change has the potential 

to adversely impact corals throughout 

the Pacific Ocean and in other parts of 

the world.

Oil and Gas Exploration and Extraction

Since 1958, production of oil and gas 

has occurred from offshore platforms 

in southern California. A total of 26 

platforms (23 in federal waters, 3 in 

state waters) are currently sited off Pt. 

Arguello, San Pedro and in the Santa 

Barbara Channel at water depths 

ranging from 11 to 363 m (Love et al. 

2003, Figures. 3.11-3.13). A permanent 

moratorium is currently in place on new 

oil and gas leases in California state 

waters, and a moratorium on new leases 

in federal waters is in affect until 2012 

(John Romero, MMS, pers. comm.). 

No restrictions are in place for existing 

lease sites, although no platforms have 

been erected since 1989 (Nevarez et al. 

1998). As of 1 May 2006, 1290 total wells had 

been drilled at the 23 platforms in federal waters. 

In 2005, 43 of 79 active leases were producing 

and 23 development wells were spudded (MMS 

2006). One potential threat to corals and other 

filter-feeding organisms from offshore oil and 

gas operations is through the introduction of 

suspended or re-suspended materials (Cimberg 

et al. 1981; Raimondi et al. 1997; NOAA 2004). 

During drilling, a mixture of water, clay, and barite 

is pumped down the well hole primarily to cool 

the equipment and dislodge any rocks. Hyland 

et al. (1994) documented the abundance of hardbottom epifauna before, during and after drilling 

began at three platforms. They found a decrease 

in the density of four invertebrate taxa, including 

Caryophyllia sp., after drilling began. 

Deployment of Gas Pipelines and 

Communication Cables 

Numerous gas pipelines connect offshore 

production platforms to shore side facilities in 

southern California (Figures. 3.11-3.13). In 

addition, cable routes transect many parts of the 

continental margin off all three west coast states. 

Recent cable installations include the Alaska 

United Fiber System – West, which extends from 

Warrenton, Oregon west-southwest over the shelf 

and is buried down to 1500 meters water depth. 

A reinstallation of a communication cable that 

connects the U.S. and Japan via the northeastern 

portion of the Olympic Coast NMS was completed 

in September 2006 (Mary Sue Brancato, Olympic 

Coast NMS, pers. comm.). Future cable laying 

operations include the NEPTUNE cable system 

– a regional cabled ocean observatory in the 

Figure 3.11. Map showing locations of oil and gas leases, platforms and associated pipelines in the Santa Maria Basin off Pt. 

Arguello, California. Image source: MMS Pacific OCS Region.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

129PACIFIC COAST

Figure 3.12. Map 

showing locations of 

oil and gas leases, 

platforms and associated pipelines in 

the Santa Barbara 

Channel off southern 

California. Image 

source: MMS Pacific 

OCS Region.

Figure 3.13. Map 

showing locations of 

oil and gas leases, 

platforms and associated pipelines off 

San Pedro, California. Image source: 

MMS Pacific OCS 

Region.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

northeast Pacific Ocean. NEPTUNE will consist 

of a 3000-km network of fiber-optic cables with 

about 20 instrumented nodes located along the 

route (Figure 3.14). Portions of the cable network 

off Vancouver Island, BC are under construction, 

with the U.S. portion to be installed later. 

There have been no reports in the region of 

communication cables or gas pipelines transecting 

areas of known coral habitat. However, this may 

be due to a lack of effort on this topic. Where 

feasible, cables and pipelines are buried down 

to deep-water depths (usually 1500 m) to avoid 

interactions with commercial fishing operations. 

If corals were located nearby, they could be 

buried or smothered by resuspended sediments. 

Where cables and pipelines transect hard-bottom 

seabed, they cannot be buried and therefore may 

cause severe impacts to corals if present in the 

area. Damage can occur along the cable or 

pipeline itself or over larger swaths by the heavy 

anchors often used during placement or repair 

(Freiwald et al. 2004). 

Sedimentation

Studies throughout the world report negative 

effects of sedimentation on corals (Dodge et al. 

1974; Dodge and Vaisnys 1977; Dodge and Lang 

1983; Hardin et al. 1994); however, some taxa 

are more vulnerable than others. For example, 

two experiments have shown that scleractinians 

have the ability to actively remove sediment 

from their polyps (Reigl 1995; Shelton 1980), 

while alcyonaceans rely solely on water motion 

and gravity (Reigl 1995). Despite this ability by 

some taxa, sedimentation is still a major threat 

to the health of coral communities throughout 

the world (Norse 1993). The U.S. Pacific coast 

is no exception. At hard-bottom sites off Pt. 

Conception, CA, Hardin et al. (1994) examined 

the spatial distribution of epifaunal assemblages 

in relation to suspended sediment flux. The 

authors discovered a negative correlation 

between sediment flux within 2 m of the seafloor 

and the coverage of filter-feeding organisms that 

include two stony corals – Lophelia pertusa and 

Desmophyllum dianthus. 

Natural sources of sediments to the continental 

margin include coastal rivers – the Columbia being 

the largest in the region. Studies of sediment 

deposition and accumulation rates have been 

conducted at sites off Oregon (Kulm et al. 1975), 

northern California (Wheatcroft et al. 1997) and 

central California (Lewis et al. 2002; Eittreim et al. 

2002). Although these studies did not specifically 

address impacts to corals or others filter feeders, 

they did provide average annual deposition rates 

over broad ranges of depth and latitude. 

Other causes of sediment resuspension include 

bottom trawling, slumps, storms and turbidity 

currents (triggered by earthquakes). Bottom 

trawling may pose the highest risk because of 

its documented interactions with corals (mostly 

pennatulaceans and gorgonians in the form of 

bycatch) and its widespread use in the region. 

Unfortunately, there is little specific information in 

the region on the effects of any of these potential 

causes. 

Pollution

The authors are aware of only one study in the 

region that specifically addressed the effects of 

pollutants on deep coral communities. Hyland 

et al. (1994) examined the effects of chemical 

contaminants from drilling discharges on the 

benthos off Pt. Arguello, CA. The authors 

found that concentrations were below toxic 

levels, suggesting that any biological changes 

to epifauna were due to increased particulate 

loading. Because of the lack of information in the 

region and throughout the world, the effects of 

pollution on deep corals remain uncertain. 

Coral Harvest

Because it retains its pigment after death, 

Stylaster californicus is popular among 

recreational collectors and has also been 

harvested commercially for sale in shell shops 

(NMSP 2005). However, commercial harvest of 

Stylasterids is now prohibited off California.

Mineral Mining

Compared to oil and gas development, mineral 

mining has the potential to adversely impact 

larger areas of seafloor habitat (Carney 2001). 

Two mining operations have been proposed 

off the U.S. Pacific coast, but neither has been 

initiated. Manganese nodules are ores containing 

minerals including manganese, cobalt, nickel 

and copper. They were originally discovered 

during the HMS Challenger voyage, although 

later discoveries of dense aggregations in the 

central north Pacific provoked economic interest 

during the Cold War (Carney 2001). Because 

of the distance and operating depths involved 

and continued availability of key minerals from 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

131PACIFIC COAST

Figure 3.14. Map showing locations of proposed 

cable routes and node locations for the NEPTUNE 

project. Image credit:: Debbie Kelley, Univ. of Washington, Center for Environmental Visualization.

terrestrial sources, mining of the nodules was 

never initiated. These dense aggregations occur 

outside the EEZ, however, so potential mining 

operations would not impact U.S. coral resources. 

The second proposed offshore mining operation 

was for polymetallic sulfides. In 1983, the U.S. 

Minerals Management Service issued a draft 

environmental impact statement for a proposed 

lease sale of a portion of the seabed on the Gorda 

Ridge (see Figure 3.2). Despite initial studies 

as to the impact of mining in the vicinity, there 

was lack of industry interest. Although it is not 

known whether corals occur on Gorda Ridge, a 

photographic study of the proposed lease site in 

1986 revealed the presence of other suspension 

feeders (Carney 2001).

Climate Change

Although oceans are moderating climate change 

by assimilating anthropogenic CO2 emissions, it 

is not without consequence to ocean chemistry. 

Uptake of atmospheric CO2 is causing declining 

pH in ocean surface waters (already 0.1 units 

lower than preindustrial values) and is predicted 

to decrease carbonate concentrations in the 

deep ocean (Orr et al. 2005). Corals use 

carbonate ions to form skeletal components. 

Two forms of calcium carbonate are common in 

the oceans: 1) aragonite, used by scleractinians 

and most stylasterids, and 2) calcite, used by 

octocorals and about 10% of stylasterids (Cairns 

and Mcintyre 1992). Aragonite is more soluble 

than calcite; therefore its depth of saturation (i.e., 

aragonite saturation horizon (ASH)) is shallower 

than that of calcite. Furthermore, the ASH in the 

north Pacific occurs at shallower water depths 

than the north Atlantic, resulting in dissolution 

rates twice as high in the upper 1000 m (Feely 

et al. 2004). Guinotte et al. (2006) hypothesize 

that the paucity of deep-sea, bioherm-building 

scleractinians in the north Pacific is a result of this 

relatively shallow (50-600 m) ASH. The authors 

also suggested that stony corals in the north 

Pacific are already living in marginal aragonite 

saturation states. Off the U.S. Pacific coast, stony 

corals are known to occur at water depths down 

to 578 meters (Cairns 1994). One of the largest 

accumulations of colonial scleractinian observed 

thus far in the region (i.e., an extensive low-lying 

mound of Lophelia pertusa at the Olympic Coast 

NMS) was at a water depth of approximately 250 

m (Brancato et al. 2007). Guinotte et al. (2006) 

predicted that the ASH off the entire U.S. Pacific 

coast would be shallower than 200 m by 2060, 

while changes in the calcite saturation horizon 

will be less pronounced (Orr et al. 2005). If 

those predictions are correct, the distributions 

of scleractinians in the region could be severely 

impacted. Stylasterids would most likely be less 

affected by changing ASH because they occur at 

shallower water depths (<183 m).

Evidence suggests that global climate change 

may pose other threats to corals. Corals are 

sessile filter feeders, most likely feeding on 

suspended organic matter that rains down 

from the surface or is transported by currents 

(Kiriakoulakis et al. 2005). Because many of the 

organisms that comprise this source of organic 

material (e.g., coccolithophores, foraminiferans, 

pteropods) use carbonate to form protective 

shells, reduced carbonate concentrations may 

impact nutrient availability for corals. In addition, 

rising atmospheric CO2 is increasing deep-sea 

water temperatures (Barnett et al. 2005) and 

altering salinities (Curry et al. 2003), which may 

in turn cause changes in thermohaline circulation 
PACIFIC COAST

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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

(Joos et al. 1999). Because corals have evolved 

in steady-state, nutrient-rich environments, they 

may be particularly susceptible to such changes 

in environmental conditions (Guinotte et al. 

2006). 

Invasive Species

NOAA continues to search for information on 

the effects invasive species might have on deep 

corals in the region. Thus far, no specific studies 

or adverse interactions have been identified.

VII. MANAGEMENT OF FISHERY 

RESOURCES AND HABITATS

The Pacific Fisheries Management Council (the 

Council) and the National Marine Fisheries Service 

(NMFS) are responsible for the management of 

fishery resources in federal waters off the U.S. 

Pacific Coast from California to Washington. 

Fisheries that occur solely within state waters 

(e.g., pink shrimp in Oregon) are regulated by 

the individual states, along with other activities 

that may have impacts to intertidal and subtidal 

habitats. Other activities occurring within five 

national marine sanctuaries and on the outer 

continental shelf are regulated by the NOAA’s 

National Marine Sanctuary Program and the 

Department of Interior’s Minerals Management 

Service (MMS), respectively. Although specific 

protections of corals have only been implemented 

recently, a number of measures adopted by these 

federal and state agencies have had the indirect 

effect of coral habitat conservation. These 

measures will be highlighted below after a brief 

synopsis of research and mapping programs in 

the region.

Mapping and Research

A multitude of seabed information exists for the 

Pacific coast region including swath bathymetric 

data and sidescan sonar imagery collected 

during large-scale mapping programs and during 

targeted geologic investigations, an extensive 

database of sediment samples collected by the 

oil industry and during submersible surveys, 

seismic reflection profiles collected by the oil 

industry and academic institutions, structural 

geologic maps created by the state of California, 

the United States Geological Survey (USGS) 

and Oregon State University, and photographic 

and video imagery collected during numerous 

submersible and ROV surveys. This information 

was recently used in the creation of a regional 

map of benthic habitats for the entire continental 

margin (i.e., intertidal out to ca. 3000 meters water 

depth) off the Pacific coast (Figure 3.15). Two 

complimentary projects were commissioned to 

inform the recent Groundfish EFH environmental 

impact statement. The Center for Habitat Studies 

at Moss Landing Marine Laboratories created the 

map for California (Greene and Bizzarro 2005), 

while the Active Tectonics and Seafloor Mapping 

Lab at Oregon State University produced the 

maps for Oregon and Washington (Goldfinger 

et al. 2005). The maps show habitat polygons 

delineating physiographic structures (e.g., shelf, 

slope and canyons) and surficial lithology (e.g., 

mud, sand and rock) at horizontal scales ranging 

from tens of meters to a kilometer. Habitat types 

were coded based on a modified version of a 

deep-water habitat characterization scheme 

developed by Greene et al. (1999). A unique 

feature of these maps is an associated layer of 

data quality that quantifies data density and ranks 

data sources for their utility to habitat mapping 

(Romsos et al. in press). From these habitat 

maps, rocky habitats are estimated to comprise 

approximately 9% (21,000 km2) of the seafloor 

area out to ca. 3000 meters water depth.

Several federal and state agencies and academic 

institutions have conducted additional targeted 

seafloor mapping projects off the Pacific coast 

to provide base maps for a variety of benthic 

habitat investigations. Study areas that have 

incorporated underwater observational platforms 

(e.g., camera sleds, submersibles, ROVs) with 

remote geophysical mapping techniques (e.g., 

sidescan and multibeam sonar, seismic profiling, 

laser line scan) include several rocky banks and 

outcrops off southern California, the Big Sur coast, 

Davidson Seamount, Monterey Canyon, Cordell 

Bank, rocky banks on the outer shelf and reefs 

along the Oregon coast, Astoria Canyon, and hard 

bottom seafloor features in the Olympic Coast 

NMS (Figure 3.16). Olympic Coast NMS is also 

continuing its deep sea habitat mapping program 

(Intelmann 2006) to guide future ROV survey 

efforts to document deep coral communities. 

Many of these investigations have resulted in the 

discoveries of deep coral communities that were 

highlighted earlier in this chapter.

To date, there has been no regular monitoring of 

deep coral habitats in the region. In 2004 and 2006, 

the Olympic Coast NMS conducted assessments 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

133PACIFIC COAST

Figure 3.15. Map of Pacific coast of the U.S., showing benthic habitats from the coastline out to approximately 3000 meters water depth. Benthic habitat characterization was modified after Greene et al. (1999) and incorporates information on seafloor lithology (e.g., sedimentary or rocky) and physiography (e.g., shelf, slope, basin, canyon wall). Maps of Washington and Oregon habitats were created the Active Tectonics and Seafloor Mapping Lab at Oregon State University. Map of California habitats was created by the Center for Habitat Studies at Moss Landing Marine Laboratories.
PACIFIC COAST

134

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Figure 3.16. Map of the Pacific coast of the U.S., highlighting growing network of study areas and some of 

the participating organizations conducting interdisciplinary studies of fish habitat (WDFW, Washington Department of Fish and Wildlife; OCNMS, Olympic Coast National Marine Sanctuary; NWFSC, Northwest Fisheries 

Science Center; SWFSC, Southwest Fisheries Science Center; PMEL, Pacific Marine Environmental Laboratory; WSUV, Washington State University, Vancouver; OSU, Oregon State University; ODFW, Oregon Department of Fish and Wildlife; CBNMS, Cordell Bank National Marine Sanctuary; MLML, Moss Landing Marine 

Laboratories; MBNMS, Monterey Bay National Marine Sanctuary; and UCSB, University of California, Santa 

Barbara). Image source: Wakefield et al. 2005, Figure 3.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

135PACIFIC COAST

of deep coral and sponge communities, but it 

is unclear how future fiscal environments will 

affect regular monitoring efforts. The Southwest 

Fisheries Science Center (SWFSC) often 

conducts submersible surveys of hard-bottom 

habitats off California. Although the primary 

focus of these surveys is groundfish habitats, they 

often occur in areas where corals are located and 

have therefore resulted in important discoveries, 

including that of the newly described black coral, 

Antipathes dendrochristos. Other surveys by the 

Northwest Fisheries Science Center (NWFSC) 

such as those utilizing AUVs are just being 

developed as a routine method for groundfish 

and may provide more routine monitoring of deep 

coral habitats coastwide.

Directed Harvest

Presently, there is no directed harvest of corals 

in the region.

Minerals Management Service

The United States Minerals Management 

Service (MMS) is responsible for the regulation 

of development and extraction of offshore 

energy resources, and they regularly conduct 

research in the Pacific outer continental shelf 

region. Mitigation of potential impacts to the 

nearby marine environment from oil and gas 

development has been a priority of MMS since oil 

production began of southern California in 1958. 

Because of the potential risks from offshore oil 

and gas operations, MMS developed anchoring 

guidelines to minimize impacts to hard-bottom 

communities. These guidelines were crafted as 

a result of numerous studies conducted before 

and after drilling projects (Mary Elaine Helix, 

MMS, pers. comm.). In addition, a long-term 

monitoring program was conducted in the late 

1980s to evaluate environmental impacts of oil 

and gas development on marine and coastal 

resources (Steinhauer and Imamura 1990). It 

was these studies that provided some of the first 

in situ observations of stony corals in the region 

including Lophelia pertusa, Desmophyllum

dianthus and Paracyathus stearnsii.

Fishery Management Councils 

The Pacific Fishery Management Council, in 

cooperation with NMFS, has implemented a 

comprehensive plan to protect EFH for groundfish 

(see PFMC 2005, 2006). The plan was developed 

in collaboration with NGOs, the fishing industry, 

and the National Marine Sanctuary Program to 

focus largely on pristine or untrawled habitat 

and biogenic habitats such as corals that are 

vulnerable to impact from human activities. The 

plan is considered comprehensive because it 

addresses impacts from the full range of human 

activities (fishing and non-fishing), and includes 

procedures for adaptation as new information 

becomes available.

Management measures to minimize adverse 

impacts from fishing include marine protected 

areas, reductions in fishing effort, and gear 

restrictions as recommended by the National 

Research Council (NRC 2002). Over 130,000 mi2

(336,700 km2) are now marine protected areas 

(MPAs) and fully protected from impacts from 

bottom trawls, with selected vulnerable habitats 

protected from all fishing gears that contact the 

bottom (Figure 3.17). The MPAs are distributed 

the length of the coast and include both federal 

and state waters. The MPAs work in concert with 

other spatial management actions taken by the 

Council such as the Rockfish Conservation Areas 

and Cowcod Conservation Areas (Figure 3.18) 

that have significantly reduced fishing effort in 

habitats important to the adult life-stage of some 

overfished species. Some of these habitats 

are hard-bottom areas that may include deep 

corals. Fishing effort has also been significantly 

reduced off the central California coast through a 

collaboration of the fishing industry and the Nature 

Conservancy. A feature of this collaboration is the 

private purchase of trawl fishing permits by the 

Nature Conservancy. Finally, gear restrictions 

implemented by the Council include coast wide 

prohibitions on gear types known to have a high 

impact on benthic habitats. Prohibited gears 

include dredge gear, beam trawl and large 

footrope bottom trawl. 

Impacts from non-fishing activities were 

addressed by the Council through the 

establishment of Habitat Areas of Particular 

Concern (HAPC) and publication of recommended 

conservation measures (PFMC 2005). NMFS is 

now using the HAPC to focus their conservation 

efforts related to non-fishing activities through 

the EFH consultation process. 

The Council and NMFS recognized that the plan 

to protect groundfish habitat is, in large part, 

based on precautionary management principles 

due to important gaps in available information. To 

accommodate the likelihood that new information 
PACIFIC COAST

136

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Figure 3.17. Map showing locations of conservation areas specified in Amendment 19 to the Pacific coast 

Groundfish fishery management plan. Conservation areas were developed in the Pacific Coast Groundfish 

Essential Fish Habitat EIS as part of the preferred alternative to minimize adverse impacts to EFH. Image 

source: PFMC 2006.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

137PACIFIC COAST

Figure 3.18. Map showing locations of some federal marine managed areas off the U.S. Pacific Coast, including national marine sanctuaries, the Cowcod Conservation Areas (CCA) and the trawl Rockfish Conservation 

Areas (RCA). The boundaries of the trawl RCA are adjusted on two-month management cycles in response 

to bycatch information on overfished species. Shoreward boundaries can be as shallow as the shoreline while 

seaward boundaries can be as deep as 250 fathoms. An area between the lines approximating the 100- and 

150-fathom isobaths (shown on map) has been permanently closed to bottom trawling since inception of the 

trawl RCA in 2002.
PACIFIC COAST

138

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Figure 3.19. Map 

showing location of the 

Channel Islands National Marine Sanctuary 

(CINMS) and its network 

of marine protected 

areas. State MPAs were 

implemented in April 

2003 while federal MPAs 

will be implemented 

in July 2007. Image 

source: Channel Islands 

NMS. 

will become available through research, the 

Council established a streamlined process to 

adapt habitat protection measures. 

In the Magnuson-Stevens Fishery Conservation 

and Management Reauthorization Act of 2006 (P.L. 

109-479), Congress directed NOAA to implement 

a Deep Sea Coral Research and Technology 

Program and allowed the designation of deep 

coral zones. The Deep Sea Coral Research 

and Technology Program will identify, locate 

and map locations of deep sea corals, monitor 

activity in locations where deep sea corals are 

known or likely to occur, conduct research and 

develop technologies to assist fishing industry 

participants in reducing interactions between 

fishing gear and deep sea corals. The Councils 

have discretionary authority to designate zones 

for the protection of deep sea corals in areas 

where deep sea corals have been identified by 

the Deep Sea Coral Research and Technology 

Program. 

National Marine Sanctuaries

In the Pacific region there exist five national 

marine sanctuaries. The Olympic Coast NMS 

is located off northern Washington, while the 

other four sanctuaries, Cordell Bank, Gulf of the 

Farallones, Monterey Bay, and Channel Islands 

are located off central and southern California 

(Figure 3.18). Except in designated marine 

reserves or conservation areas, the sanctuaries 

do not regulate fishing including the use of bottom 

contact gears. All sanctuaries, however, prohibit 

other activities that may be harmful to corals, 

including but not limited to 1) new oil, gas or mineral 

exploration, development and production, 2) 

discharge of materials or substances except fish 

parts, bait, water or other biodegradable effluents, 

and 3) alteration of the seabed except for normal 

fishing activities and anchoring. In addition, the 

Cordell Bank NMS prohibits the removal or injury 

of benthic organisms in waters above 50 fathoms 

(91 m), except during normal fishing operations 

outside of designated conservation areas. 

In May 2007, the National Marine Sanctuary 

Program established a network of marine reserves 

and one conservation area within the Channel 

Islands NMS (Figure 3.19) to complement an 

existing network of state reserves implemented 

in April 2003 by the state of California (Figure 

3.19). After implementation in July 2007, the new 

federal reserves and conservation area will total 

110.5 nmi2 and 1.7 nmi2, respectively. The state 

of California is expected to extend the boundaries 

of it’s existing reserves to the three mile statefederal waters boundary, thereby increasing to 

total area of reserves and conservation areas to 

232 nmi2 and 8.6 nmi2, respectively. Like existing 

state reserves, all extractive activities will be 

prohibited while certain fishing for lobster and 

recreational take of pelagic fishes will be allowed 

in conservation areas (15 FR 29208). Proposed 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

139PACIFIC COAST

rules for marine zoning in the Monterey Bay and 

Cordell Bank NMS are expected in the near 

future.

State activities

The state of California in recent years has enacted 

a series of laws to direct the management of 

state marine resources, including the Marine 

Life Management Act (Stats. 1998, ch. 1052), 

the Marine Life Protection Act (MLPA, Stats. 

1999, ch. 1015), and the Marine Managed 

Areas Improvement Act (Stats. 2000, ch. 385). 

In particular, a key mandate of the MLPA is to 

design and manage a network of MPAs to 

protect marine life, habitats, ecosystems, and 

natural heritage while providing recreational, 

educational and study opportunities. In addition 

to the marine protected areas implemented in 

2003, the state, in April 2007 adopted a network 

of 29 MPAs for their Central Coast Study Region, 

which encompasses state waters between Pt. 

Conception and Pigeon Point (Figure 3.20). 

These include new and expansions of existing 

MPAs that cover approximately 204 mi2 (528 

km2) or 18% of state waters in the central coast 

region, including 97 mi2 (251 km2) of no-take 

reserves (Office of Administrative Law Notice File 

Number Z06-1031-05). While corals (specifically 

hydrocorals) are listed as “benefit species” for 

only three of the MPAs, the protection of both 

shallow and deep hard bottom habitats is an 

objective for this network of MPAs. Planning 

and scoping of additional MPAs began in early 

2007 for the next study region, the North Central 

Coast, which encompasses state waters from 

Pigeon Point north to about Pt. Arena.

To date, only about a third of the seafloor in 

California state waters has been mapped at any 

appreciable scale (Rikk Kvitek, CSUMB, pers. 

comm.). The Seafloor Mapping Lab of California 

State University Monterey Bay (CSUMB) and 

its partners are currently engaged in the North 

Central Coast Mapping Project, which will produce 

multibeam base maps for state waters seaward 

of the 20-meter isobath from Año Nuevo north to 

Pt. Arena. Another project mapping the coastal 

area in the Santa Barbara Channel involves the 

Seafloor Mapping Lab, USGS and the California 

Ocean Protection Council. Over the next 5-6 

years, the California Coastal Conservancy and 

Ocean Protection Council hope to complete 

comprehensive mapping for the remaining 66% of 

state waters (Rikk Kvitek, CSUMB, pers. comm.). 

The time frame for completion will depend on 

available funds.

Oregon also suffers from lack of seabed 

information in state waters. To date, less than 

5% of Oregon’s ca. 950 nm2 (3263 km2) territorial 

sea is mapped. In March 2006, a consensus 

statement, signed by 20 Oregon marine scientists, 

called for support and funds to map the seafloor 

within Oregon’s territorial sea. Projected costs 

are under $6 million. Stated reasons for this 

mapping plan include management of hazards 

posed by tsunami events, describing nearshore 

habitats on which nearshore fisheries and 

marine resources depend, and scientific support 

of two gubernatorial proposals – establishment 

of a limited network of marine reserves in state 

waters, and a national marine sanctuary to be 

sited off Oregon. In response to the scientific 

consensus statement, the Territorial Sea Mapping 

Bill (HB 2924), which would appropriate funds for 

seafloor mapping, was introduced in February 

2007. Passage of HB 2924 along with the 

establishment of reserves and a national marine 

sanctuary are still pending.

In September 2006, the three west coast governors 

signed a joint agreement to collaborate on critical 

ocean and coastal protection and management 

issues. Short-term priorities for this collaboration 

include increased funding for mitigation of 

nonpoint source pollution, opposition to oil and 

gas leasing, exploration and development, and a 

regional research plan to support ocean observing 

programs and seafloor and habitat mapping.

VIII. REGIONAL PRIORITIES TO 

UNDERSTAND AND CONSERVE DEEP 

CORAL COMMUNITIES

Given the limitations of existing information off the 

U.S. Pacific coast, it is clear that more targeted 

data collections and mapping efforts are needed. 

Because many collections are made from long 

trawls that can traverse several habitats, it is 

impossible to determine specifically the habitat 

from which these species were collected. 

Therefore, to date, it is difficult to map corals at 

a regional scale showing the appropriate habitat 

associations. Furthermore, coral specimens are 

continuously collected during in situ photographic 

surveys, regional trawl surveys and by fisheries 

observers. Rapid identification of these samples is 
PACIFIC COAST

140

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Figure 3.20. Map showing locations of marine protected areas in the Central Coast study region for the state 

of California, including state marine conservation areas (SMCA), marine parks (SMP), marine reserves (SMR) 

and marine recreational management areas (SMRMA). Image source: California Department of Fish and 

Game.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

141PACIFIC COAST

needed to mitigate further impacts to deep corals 

and their habitats. Finally, the establishment of 

a network of conservation areas in the region 

provides an unprecedented opportunity to 

monitor coast wide recovery of benthic habitats 

(including those that support coral communities) 

from fishing impacts. 

Mapping

Because little is known about the nature of 

relationships between corals, other invertebrates 

and demersal fishes off the Pacific coast, the 

highest mapping priority is to quantify those 

relationships. To date, few studies (see Tissot 

et al. 2006; Pirtle 2005; Hardin et al. 1994) 

in the region have examined the nature of 

relationships between corals, other structureforming invertebrates and fishes, though analysis 

of recent surveys is ongoing (see Brancato et al. 

2007). In order to evaluate the importance of 

corals to their benthic communities, future in situ

surveys will need to incorporate a more holistic 

investigation of species relationships and habitat 

characteristics.

Surveys are needed that specifically monitor 

the abundance and distribution of corals in 

representative habitat types so that accurate 

and comprehensive maps of their distribution 

and abundance can be made. Targeted surveys 

on representative habitats using underwater 

platforms (e.g., submersibles, ROVs, AUVs) in 

conjunction with optical survey systems (e.g., 

video, laser line scan) are needed. Survey areas 

can be selected using maps of surficial geologic 

habitats that were developed for the Pacific coast 

Groundfish EFH EIS (see Figure 3.15) or higher 

resolution maps of specific areas. In addition, 

areas of repeated coral bycatch identified during 

mapping of existing observer and trawl survey 

data, should be intensively mapped to establish 

the extent of these “hot spots.” 

Research

Coral information can be collected ancillary to 

other survey and monitoring activities, however 

it is difficult for non-specialists on these surveys 

to provide specific identifications. Collaborations 

with systematics experts at museums through 

a pilot project in the NWFSC’s Genetics and 

Evolution Program to develop molecular methods 

for the rapid identification of Pacific coast 

corals are already in place. The development 

of this capability will provide for species-level 

identification of corals and potentially other 

difficult to identify structure-forming invertebrates 

from the region’s ongoing trawl surveys and 

observer program and will provide validations 

of identifications during targeted deep coral 

surveys. 

Deep corals are vulnerable to a variety of 

activities including oil and gas exploration and 

fishing; however, little is known about the potential 

recovery rates of these species. There are a 

variety of ways to monitor recovery. For instance, 

age and growth information can be obtained 

through a combination of biochemical studies, 

14C and other radiometric dating, stable oxygen 

isotope data from the corals themselves and 

other biochemical analyses. In addition to basic 

age information, it should be possible to associate 

growth stages of corals to environmental change. 

Working with biochemists at collaborating 

academic institutions, the physiological potential 

for recovery of deep corals via geochemical and 

biochemical analyses of age and growth as well 

as growth potential can be investigated.

Other gaps in information result from the fact 

that few targeted surveys have been conducted 

in the region. Nearly thirty species of stylasterid 

corals have been observed in waters off Alaska 

including the Aleutian Islands coral gardens and 

parts of the Bering Sea and Gulf of Alaska (Stone 

and Shotwell, Chapter 2). In contrast, only five 

species of stylasterids have been observed 

in the Oregon Province with an additional two 

species found off British Columbia (Appendix 

3.1, Jamieson et al. 2006). This may be a result 

of sampling bias; nonetheless, more focus needs 

to be given to the shallower depths where these 

taxa are commonly found. Also, reef-building 

scleractinians (e.g., Lophelia pertusa) while 

common in the San Diego Province do not form 

expansive reefs like those in the north Atlantic. 

While ocean chemistry may be a factor (see 

Guinotte et al. 2006; Orr et al. 2005), it’s clear, 

especially with the recent discovery of a Lophelia 

mound in the Olympic Coast NMS that more 

attention needs to be given to these taxa. 

Although more research on deep coral 

communities is needed, data mining opportunities 

do exist. Since the late 1980s, underwater vehicles 

and camera sleds have been used to survey 

the benthic environment off the Pacific coast. A 

product of these surveys is hours upon hours of 
PACIFIC COAST

142

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

video data. Review of these videos, while timeconsuming, may provide additional observations 

of deep corals and insights into their ecology 

(see Strom 2006). A great example of rescuing 

existing data from past visual surveys of deep 

corals and other structure-forming invertebrates 

is a longterm and productive collaboration among 

fishery biologists at the SWFSC and invertebrate 

ecologists at Washington State University. This 

collaboration has resulted in a georeferenced 

database on the distribution and abundance of 

deep corals and associated fishes and habitats off 

California, as well as providing the informational 

basis for evaluating deep corals as EFH for 

demersal fishes. 

Summary of research priorities:

• Quantify and describe the nature of 

associations of deep corals with other 

structure-forming invertebrates and demersal 

fishes.

• Timely identification of coral specimens 

collected during region trawl surveys and by 

fisheries observers.

• Investigate recovery rates of corals and 

their habitats from a variety of stressors, 

particularly fishing.

• Focus more survey effort in shallow rocky 

habitats where corals may be more susceptible 

to coastal activities.

• Data mining of video records from numerous 

submersible and ROV surveys of rocky 

features throughout the region to identify 

deep corals and any significant habitat and 

species associations.

IX. CONCLUSION

The Pacific coast from California to Washington 

hosts a considerable amount of deep coral 

habitat. Significant coral communities have been 

discovered at Davidson Seamount (DeVogelaere 

2005), in Monterey Canyon, at Cordell Bank (Pirtle 

2005), in the Olympic Coast NMS (Brancato et al. 

2007) and on numerous rocky banks off southern 

California (Love et al. 2007; Tissot et al. 2006; 

Yoklavich and Love 2005), including the recently 

discovered Christmas tree coral, Antipathes 

dendrochristos (Opresko 2005). Pennatulaceans, 

black corals and gorgonians have also been 

recorded coast wide in the catch of bottom trawl 

surveys and by fishery observers. For the most 

part, corals in the region do not build reefs with 

observations of only two reef-building stony 

corals - Lophelia pertusa and Madrepora oculata

in the San Diego Province and L. pertusa in the 

Olympic Coast NMS. Although associations of 

corals with other invertebrates and fishes have 

been reported, there is no direct evidence that any 

of these represent obligate relationships between 

taxa. More targeted studies are needed to further 

investigate and quantify these relationships.

Much of the recent information on the regional 

zoogeography of higher-level coral taxa was 

collected during bottom trawl surveys. Some 

of these surveys are ongoing and will provide 

continued mapping information within the 

limitations of the collection methods on these 

species. More detailed information, but in a 

limited geographic scope, has been collected 

using submersibles, remotely operated vehicles 

(ROVs) and more recently, autonomous 

underwater vehicles (AUVs); however, these 

surveys most often focus on demersal fishes, 

though usually in areas where deep corals 

are found. Unlike trawl surveys, these in situ

photographic surveys can provide information 

on the relationships between deep corals, other 

invertebrates and demersal fishes. In situ

photographic surveys provide localized mapping 

of deep coral habitats but are seldom conducted 

on a repeated basis, and therefore do not provide 

routine monitoring of these sites. Additional 

information on the distribution of corals as well 

as monitoring fishing impacts can be gleaned 

from information collected by fisheries observers. 

Observers currently collect this information on an 

ongoing basis. 

Unique attributes of the region that may influence 

coral distributions include a narrow continental 

shelf, coastal upwelling and a large latitudinal 

extent encompassing two distinct zoogeographic 

provinces. Unlike the eastern coast of the 

U.S., the Pacific coast is part of an active 

continental margin, with both prolonged (e.g., 

subduction) and episodic (e.g., turbidity currents) 

events that both shape and impact deep coral 

habitats. However, the most significant and 

immediate threat to coral habitats in the region 

is commercial fishing, namely widespread 

use of bottom trawls to target demersal fishes. 

Direct evidence of fishing impacts to corals has 

been reported by fishery observers. Additional 

evidence of fishery interactions with the seafloor 

(e.g., trawl marks in sediments, derelict gear) in 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

143PACIFIC COAST

the vicinity of corals also is provided from in situ

photographic surveys, such as those conducted 

recently in the Olympic Coast NMS (Hyland et al. 

2004; Brancato et al. 2007). With recent fishery 

management measures (e.g., area closures, gear 

restrictions), however, the risk posed by bottom 

trawling has been significantly reduced. Although 

conservation areas encompass many known coral 

habitats coastwide, recent discoveries of diverse 

coral communities outside of conservation areas 

(e.g., in the Olympic Coast NMS) warrant further 

consideration of protection.

X. REFERENCES

Allen MJ, Groce AK, Diener D, Brown J, Steinert 

SA, Deets G, Noblet JA, Moore SL, Diehl 

D, Jarvis ET, Raco-Rands V, Thomas C, 

Ralph Y, Gartman R, Cadien D, Weisberg 

SB, Mikel T (2002) Southern California 

Bight 1998 Regional Monitoring Program: 

V. demersal fishes and megabenthic 

invertebrates. Southern California Coastal 

Water Research Project, Westminster, 

California

Allen MJ, Moore SL, Schiff KC, Weisberg SB, 

Diener D, Stull JK, Groce A, Mubarak J, 

Tang CL, Gartman R (1998) Southern 

California Bight 1994 Pilot Project: 

V. Demersal fishes and megabenthic 

invertebrates. Southern California Coastal 

Water Research Project, Westminster, 

California

Andrews AH, Cailliet GM, Kerr LA, Coale KH, 

Lundstrom C, DeVogelaere A (2005) 

Investigations of age and growth for 

three deep-sea corals from the Davidson 

Seamount off central California. Pages 

1021-1038 in Freiwald A, Roberts JM 

(eds.), Cold-water corals and ecosystems. 

Springer-Verlag Berlin Heidelberg

Aspen Environmental Group (Aspen) (2006) 

Tranquillon ridge oil and gas development 

project: Environmental impact report, 

State clearinghouse no. 2006021055, 

Prepared for County of Santa Barbara 

Planning and Development

Auster PJ (2005) Are deep-water corals important 

habitat for fishes? Pages 747-760 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin Heidelberg

Barnett TP, Pierce DW, AchutaRao K, Gleckler 

PJ, Santer BD, Gregory JM, Washington 

WM (2005) Penetration of human-induced 

warming into the world’s oceans. Science 

309:284–287
PACIFIC COAST

144

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Batchelder HP, Barth JA, Kosro PM, Strub 

PT, Brodeur RD, Peterson WT, Tynan 

CT, Ohman MD, Botsford LW, Powell 

TM, Schwing FB, Ainley DG, Mackas 

DL, Hickey BM, Ramp SR (2002) The 

GLOBEC Northeast Pacific California 

Current System program. Oceanography 

15:36-47

Bellman MA, Heppell SA, Goldfinger C (2005) 

Evaluation of a U.S. west coast groundfish 

habitat conservation regulation via 

analysis of spatial and temporal patterns 

of trawl fishing effort.Canadian Journal of 

Fisheries and Aquatic Sciences 62:2886-

2900

Brancato MS, Bowlby CE, Hyland J, Intelmann 

SS, Brenkman K (2007) Observations of 

deep coral and sponge assemblages in 

Olympic Coast National Marine Sanctuary, 

Washington. Cruise Report: NOAA Ship 

McArthur II Cruise AR06-06/07, 46pp

Briggs JC (1974) Marine Zoogeography. McGrawHill Book Company. New York, 475pp

Brodeur RD (2001) Habitat-specific distribution of 

Pacific ocean perch (Sebastes alutus) in 

Pribilof Canyon, Bering Sea. Continental 

Shelf Research 21:207-224

Cairns SD (1994) Scleractinia of the temperate 

North Pacific. Smithsonian Contributions 

to Zoology No. 557

Cairns SD (1983) A generic revision of the 

Stylasterina (Coelenterata: Hydrozoa), 

Part 1, Description of the genera. Bulletin 

of Marine Science 33:427–508

Cairns SD, Bayer FM (2005) A review of the genus 

Primnoa (Octocorallia: Gorgonacea: 

Primnoidae), with the description of two 

new species. Bulletin of Marine Science 

77:225-256

Cairns SD, Stanley GD (1982) Ahermatypic coral 

banks: Living and fossil counterparts. 

Proceedings of the Fourth International 

Coral Reef Symposium: 1:611-613

Carey Jr. AG, Taghon GL, Stein DL, Rona PA 

(1990) Distributional ecology of benthic 

megaepifauna and fishes in the Gorda 

Ridge Axial Valley. Pages 225-240 In: 

McMurray GR (ed) Gorda Ridge: A 

seafloor spreading center in the United 

States’ exclusive economic zone. 

Springer, New York

Carney RS (2001) Management applicability 

of contemporary deep-sea ecology and 

reevaluation of Gulf of Mexico studies. 

Final Report. OCS Study MMS 2001-

095. U.S. Dept. of the Interior Minerals 

Management Service, New Orleans, 

Louisiana, USA, 174pp

Cimberg RL, Gerrodette T, Muzik K (1981) Habitat 

requirements and expected distribution of 

Alaska coral. Final Report, Research Unit 

No. 601, U.S. Office of Marine Pollution 

Assessment, Alaska Office, 54pp

Clague, DA, Zierenberg R, Davis A, Goffredi S, 

McClain J, Maher N, Olsen E, Orphan V, 

Ross S, Von Damm K (2001) MBARI’s 

2000 expedition to the Gorda Ridge. 

RIDGE Events, 11:5–12

Clague DA, Holmes ML (1987). Geology, 

petrology, and mineral potential of the 

Gorda Ridge. In Scholl DW, Grantz A, 

Vedder JG (eds.) Geology and resource 

potential of the continental margin of 

western North America and adjacent 

ocean basins—Beaufort Sea to Baja 

California. Circum-Pacific Council for 

Energy and Mineral Resources, Earth 

Science Series, Houston, Texas, USA, 

pp563-580

Curry R, Dickson B, Yashayaev I (2003) A change 

in the freshwater balance of the Atlantic 

Ocean over the past four decades. Nature 

426:826–829

Dall WA (1884) On some Hydrocorallinae from 

Alaska and California. Proceedings of the 

Biological Society of Washington 2:111-

115
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

145PACIFIC COAST

Davis EE, Karsten JL (1986) On the cause 

of the asymmetric distribution of 

seamounts about the Juan de Fuca 

Ridge: Ridge crest migration over a 

heterogeneous asthenosphere. Earth and 

Planetary Science Letters 79:385-396

Department of Fisheries and Oceans Canada 

(DFO) (2000) Hexactinellid sponge reefs 

on the British Columbia continental shelf: 

geological and biological structure. DFO 

Pacific Region Habitat Status Report 

2000/02

Deuser WG (1975) Reducing environments. 

In Riley JP, Skirrow G (eds) Chemical 

Oceanography, 2nd ed., vol 3. Academic 

Press, pp1-37

DeVogelaere AP, Burton EJ, Trejo T, King CE, 

Clague DA, Tamburri MN, Cailliet GM, 

Kochevar RE, Douros WJ (2005) Deepsea corals and resource protection at 

the Davidson Seamount, California, 

U.S.A. Pages 1189-1198 in Freiwald A, 

Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Dodge R, Aller R, Thomsom J (1974) Coral 

growth related to resuspension of bottom 

sediments. Nature 247:574-577

Dodge R, Lang J (1983) Environmental correlates 

of hermatypic coral ( Montastrea annularis) 

growth on the East Flower Gardens Bank, 

north west Gulf of Mexico. Limnology and 

Oceanography 28:228-240

Dodge R, Vaisnys J (1977) Coral populations 

and growth patterns: responses to 

sedimentation and turbidity associated 

with dredging. Journal of Marine Research 

35:715-730

Eittreim SL, Xu JP, Noble M, Edwards BD (2002) 

Towards a sediment budget for the Santa 

Cruz shelf. Marine Geology 181:235-248

Etnoyer P, Morgan L (2003) Occurrences of 

habitat-forming deep sea corals in the 

Northeast Pacific Ocean. A report to 

NOAA’s Office of Habitat Conservation. 

Marine Conservation Biology Institute, 

Redmond, Washington, USA, 33pp

Feely RA, Sabine CL, Lee K, Berelson W, Kleypas 

J, Fabry VJ, Millero FJ (2004) Impact of 

anthropogenic CO2 on the CaCO3 system 

in the oceans. Science 305:362–366

Fisher WK (1938) Hydrocorals of the North Pacific 

Ocean. Proceedings of the United States 

National Museum 84:493-554

Fisk MR, Duncan RA, Fox CG, Witter JB 

(1993) Emergence and petrology of 

the Mendocino Ridge. Marine and 

Geophysical Research 15:283-296

Freiwald A, Fosså JH, Grehan A, Koslow T, 

Roberts JM (2004) Cold-water coral reefs. 

UNEP-WCMC, Cambridge, UK, 85pp

Fosså JH, Mortensen PB, Furevik DM (2002) 

The deep-water coral Lophelia pertusa in 

Norwegian waters: distribution and fishery 

impacts. Hydrobiologia 471:1-12 

Greene HG, Bizzarro JJ (2005) Essential fish 

habitat characterization and mapping 

of the California continental margin. In

Risk assessment for the Pacific coast 

groundfish FMP. Prepared for the Pacific 

States Marine Fisheries Commission, 

Portland, Oregon, USA, Appendix 3

Greene HG, Yoklavich MM, Starr RM, O’Connell 

VM, Wakefield WW, Sullivan DE, McRea 

Jr JE, Cailliet GM (1999) A classification 

scheme for deep seafloor habitats. 

Marine benthic habitats and their living 

resources: monitoring, management and 

applications to Pacific Island countries. 

Oceanologica Acta 22:663-678
PACIFIC COAST

146

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Goldfinger C, Romsos C, Robison R, Milstein R, 

Myers B (2005) Interim seafloor lithology 

maps for Oregon and Washington, 

Version 1.0. In Risk assessment for the 

Pacific coast groundfish FMP. Prepared 

for the Pacific States Marine Fisheries 

Commission, Portland, Oregon, USA, 

Appendix 2

Guinotte JM, Orr J, Cairns S, Freiwald A, Morgan 

L, George R (2006) Will human-induced 

changes in seawater chemistry alter the 

distribution of deep-sea scleractinian 

corals? Frontiers in Ecology and 

Environment 4:141-146

Hardin DD, Toal J, Parr T, Wilde P, Dorsey K (1994) 

Spatial variation in hard-bottom epifauna 

in the Santa Maria Basin, California: the 

importance of physical factors. Marine 

Environmental Research 37:165-193

Hickey BM (1998) Coastal oceanography of 

western North America from the tip of 

Baja California to Vancouver Island. In: 

Robinson AR, Brink KH (eds.) The Sea, 

Vol. 11. John Wiley & Sons, Inc, New 

York, pp345-393

Hickey BM (1979) The California current system 

– hypotheses and facts. Progress in 

Oceanography 8:191-279

Hixon MA, Tissot BN, Pearcy WG (1991) Fish 

assemblages of rocky banks of the 

Pacific Northwest [Heceta, Coquille, and 

Daisy Banks], Final Report. OCS Study 

MMS 91-0052, U.S. Dept. of the Interior 

Minerals Management Service, Camarillo, 

California, USA, 410pp

Hyland J, Cooksey C, Bowlby E, Brancato MS, 

Intelmann S. (2005) A pilot survey of 

deepwater coral/sponge assemblages 

and their susceptibility to fishing/harvest 

impacts at the Olympic Coast National 

Marine Sanctuary (OCNMS). Cruise 

Report for NOAA Ship McARTHUR II 

Cruise AR-04-04: Leg 2. NOAA Tech. 

Memo. NOS NCCOS 15. NOAA/NOS 

Center for Coastal Environmental Health 

and Biomolecular Research, Charleston, 

SC, USA, 13pp

Hyland J, Hardin D, Steinhauer M, Coats D, 

Green R, Jeff J (1994) Environmental 

impact of offshore oil development on 

the outer continental shelf and slope 

off Point Arguello, California. Marine 

Environmental Research 37:195-229

Intelmann SS (2006) Habitat Mapping Effort 

at the Olympic Coast National Marine 

Sanctuary – Current Status and Future 

Needs. Marine Sanctuaries Conservation 

Series NMSP-06-11. U.S. Department 

of Commerce, National Oceanic and 

Atmospheric Administration, National 

Marine Sanctuary Program, Silver Spring, 

MD. 29 pp

Jamieson GS, Pellegrin N, Jessen S (2006) 

Taxonomy and zoogeography of cold 

water corals in explored areas of coastal 

British Columbia. Research Document 

2006/062, Fisheries and Oceans Canada, 

45pp

Johnson KA (2002) A review of national and 

international literature on the effects 

of fishing on benthic habitats. NOAA 

Technical Memorandum NMFS-F/SPO57, 72 pp

Joos F, Plattner GK, Stocker TF, Marchal O, 

Schmittner A (1999) Global warming and 

marine carbon cycle feedbacks on future 

atmospheric CO2. Science 128:464-467

Kaiser MJ (2004) Fish in deep-water coral 

habitats. Letters to the Editor. Science 

304:1595

Kiriakoulakis K, Harper E, Wolff GA, Freiwald 

A, Grehan A, Roberts JM (2005) Lipids 

and nitrogen isotopes of two deep-water 

corals from the North-East Atlantic: initial 

results and implications for their nutrition. 

Pages 715-729 in Freiwald A, Roberts JM 

(eds.), Cold-water corals and ecosystems. 

Springer-Verlag Berlin Heidelberg
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

147PACIFIC COAST

Koenig CC, Shepard AN, Reed JK, Coleman 

FC, Brooke SD, Brusher J, Scanlon KM 

(2005) Habitat and fish populations in the 

deep-sea Oculina coral ecosystem of the 

western Atlantic. In: Barnes PW, Thomas 

JP (eds.) Benthic habitats and the effects 

of fishing. American Fisheries Society, 

Bethesda, Maryland, USA, pp795-806

Krieger KJ, Wing BL (2002) Megafaunal 

associations with deepwater corals 

(Primnoa sp.) in the Gulf of Alaska. 

Hydrobiologica 471:83-90

Kulm LD, Roush RC, Harlett JC, Neudeck 

RH, Chambers DM, Runge EJ (1975) 

Oregon continental shelf sedimentation: 

Interrelationships of facies distribution 

and sedimentary processes. Geology 

83:145-175

Lewis RC, Coale KH, Edwards BD, NowickiDouglas J, Burton EJ (2002) Accumulation 

rate and mixing of shelf sediments in the 

Monterey Bay National Marine Sanctuary. 

Marine Geology 181:157-169

Love MS, Yoklavich MM, Black BA, Andrews AH 

(2007) Age of black coral (Antipathes 

dendrochristos Opresko, 2005) colonies, 

with notes on associated invertebrate 

species. Bulletin of Marine Science. 

80:391-400

Love MS, Schroeder DM, Nishimoto MM 

(2003) The ecological role of oil and 

gas production platforms and natural 

outcrops on fishes in southern and central 

California: a synthesis of information. 

U.S. Dept. of Interior U.S. Geological 

Survey, Biological Resources Division, 

Seattle, Washington, USA, OCS Study 

MMS 2003-032

Minerals Management Service (MMS) (2006) 

Current facts and figures in the Pacific 

OCS region as of May 1, 2006. U.S. 

Minerals Management Service, Online:

http://www.mms.gov/omm/pacific/

offshore/currentfacts.htm

Morgan LE, Chuendpagdee R (2003) Shifing 

Gears: addressing the collateral impacts 

of fishing methods in U.S. Waters. Pew 

Science Series, Washington DC, 42pp

National Marine Fisheries Service (NMFS) 

(2007, in prep) Our living oceans. The 

status of habitats of U.S. living marine 

resources, 2006. U.S. Dep. Commer., 

NOAA Technical Memorandum NMFS-F/

SPO-75

National Marine Sanctuary Program (NMSP) 

(2005) Biological and historical/cultural 

resources of the study area for the Channel 

Islands National Marine Sanctuary 

management plan update. In U.S. Dept. 

of Commerce. National Oceanic and 

Atmospheric Administration. National 

Marine Sanctuary Program. Channel 

Islands National Marine Sanctuary Draft 

Management Plan / Draft Environmental 

Impact Statement. Silver Spring, MD.

National Research Council (NRC) (2002) Effects 

of trawling and dredging on seafloor 

habitat. National Academy Press, 

Washington D.C.

Nevarez L, Molotch H, Shapiro P, Bergstrom R 

(1998) Petroleum extraction in Santa 

Barbara County, California: an industrial 

history. U.S. Dept. of Interior Minerals 

Management Service, OCS Study 

MMS98-0048

Norse, EA (1993) Global marine biological 

diversity. A strategy for building 

conservation into decision making. Island 

Press, Washington DC, 383pp

Northwest Fisheries Science Center (NWFSC) 

(2007) Data report and summary analyses 

of open access fixed-gear fisheries in 

waters less than 50 fathoms. Northwest 

Fisheries Science Center, Seattle, 

Washington, USA

Northwest Fisheries Science Center (NWFSC) 

(2006) West coast groundfish observer 

program data report and summary 

analyses of limited-entry trawl permits. 

Northwest Fisheries Science Center, 

Seattle, Washington, USA
PACIFIC COAST

148

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

Opresko DM (2005) A new species of 

antipatharian coral (Cnidaria: Anthozoa: 

Antipatharia) from the southern California 

bight. Zootaxa 852:1-10

Opresko DM (2003) Revision of the 

Antipatharia (Cnidaria: Anthozoa). 

Part III. Cladopathidae. Zoologische 

Mededelingen (Leiden) 77:495-536.

Opresko DM (2002) Revision of the 

Antipatharia (Cnidaria: Anthozoa). 

Part II. Schizopathidae. Zoologische 

Mededelingen (Leiden) 76:411-442.

Orr JC, Fabry VJ, Aumont O, Bopp L, Doney 

SC, Feely RA, Gnanadesikan A, Gruber 

N, Ishida A, Joos F, Key RM, Lindsay 

K, Maier-Reimer E, Matear R, Monfray 

P, Mouchet A, Najjar RG, Plattner GK, 

Rodgers KB, Sabine CL, Sarmiento 

JL, Schlitzer R, Slater RD, Totterdell IJ, 

Weirig MF, Yamanaka Y, Yool A (2005) 

Anthropogenic ocean acidification over 

the twenty-first century and its impact on 

calcifying organisms. Nature 437:81–86.

Ostarello GL (1973) Natural history of the 

hydrocoral Allopora californica (Verrill, 

1866). The Biological Bulletin 145:548–

564

Pacific Fishery Management Council (PFMC) 

(2006) Amendment 19 to the Pacific coast 

groundfish fishery management plan: 

Essential fish habitat designation and 

minimization of adverse impacts. Pacific 

Fishery Management Council, Portland, 

Oregon, USA

Pacific Fishery Management Council (PFMC) 

(2005) Pacific coast groundfish fishery 

management plan: essential fish habitat 

designation and minimization of adverse 

impacts: Final environmental impact 

statement. Pacific Fishery Management 

Council, Portland, Oregon, USA

Pacific Fishery Management Council (PFMC) 

(2003) Fishery management plan and 

environmental impact statement for US 

west coast for highly migratory species. 

Pacific Fishery Management Council, 

Portland, Oregon, USA, 756pp

Parrish FA (2004) Foraging landscape of the 

Hawaiian monk seal. Ph.D diss., Univ. 

Hawaii, Manoa, USA, 146 pp

Pearcy WG, Stein DL, Hixon MA, Pikitch EK, 

Barss WH, Starr RM (1989) Submersible 

observations of deep-reef fishes of Heceta 

Bank. Fishery Bulletin 87:955-965

Pierce SD, Smith RL, Kosro PM, Barth JA, Wilson 

CD (2000) Continuity of the poleward 

undercurrent along the eastern boundary 

of the mid-latitude North Pacific. DeepSea Research II 47:811–829

Pirtle JL (2005) Habitat-based assessment 

of structure-forming megafaunal 

invertebrates and fishes on Cordell Bank, 

California. MS Thesis, Washington State 

Univ., Vancouver, Washington, USA, 

64pp

Raimondi PT, Barnett AM, Krause PR (1997) 

The effects of drilling muds on marine 

invertebrate larvae and adults. 

Environmental Toxicology and Chemistry 

16:1218-1228

Riegl B (1995) Effects of sand deposition on 

scleractinian and alcyonacean corals. 

Marine Biology 121:517-526.

Romsos CG, Goldfinger C, Robison R, Milstein 

RL, Chaytor JD, Wakefield WW (in press) 

Development of a regional seafloor 

surficial geologic habitat map for the 

continental margins of Oregon and 

Washington, USA. In Todd BJ, Greene 

HG (eds) Mapping the Seafloor for Habitat 

Characterization. Geological Association 

of Canada, Special Paper 47

Scott PV, Blake JA (eds.) (1998) Taxonomic 

atlas of the benthic fauna of the Santa 

Maria Basin and western Santa Barbara 

Channel. Volume 3. Special Publications 

of the Santa Barbara Museum of Natural 

History, Santa Barbara, CA, USA

Shelton GAB (1980) Lophelia pertusa (L.): 

electrical conduction and behaviour in a 

deep-water coral. Journal of the Marine 

Biological Association of the United 

Kingdom 60: pp517-528
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

149PACIFIC COAST

Smith CR, Hamilton SC (1983) Epibenthic 

megafauna of a bathyal basin off southern 

California: patterns of abundance, 

biomass, and dispersion. Deep-Sea 

Research 30:907-928

Southern California Association of Marine 

Invertebrate Taxonomists (SCAMIT) 

(2001) A taxonomic listing of soft bottom 

macro- and megainvertebrates from 

infaunal and epibenthic monitoring 

programs in the Southern California 

Bight. Ed. 4. San Pedro, California, USA, 

192pp

Stein DL, Tissot BN, Hixon MA, Barss WH (1992) 

Fish-habitat association on a deep reef at 

the edge of the Oregon continental shelf. 

Fishery Bulletin 90:540-551

Steinhauer M, Imamura E (1990) California OCS 

phase II monitoring program: Year three 

annual report: Volume I. U.S. Dept. of 

Interior Minerals Management Service, 

Pacific OCS Region, Los Angeles

Stone RP (2006) Coral habitat in the Aleutian 

Islands of Alaska: depth distribution, finescale species associations, and fisheries 

interactions. Coral Reefs 25:229-238

Stone RP, Masuda MM, Malecha PW (2005) 

Effects of bottom trawling on soft-sediment 

epibenthic communities in the Gulf of 

Alaska. In: Barnes PW, James JP (eds.) 

Benthic habitats and the effects of fishing. 

American Fisheries Society Symposium 

41, Bethesda, Maryland, USA, pp461-

475

Strom NA (2006) Structure-forming benthic 

invertebrates: habitat distributions on 

the continental margin of Oregon and 

Washington. MS Thesis, Oregon State 

Univ., Corvallis, Oregon, USA, 96pp

Strub PT, James C (2000) Altimeter-derived 

variability of surface velocities in the 

California Current System: 2. Sea­sonal 

circulation and eddy statistics. Deep-Sea 

Research II 47:831-870

Syms C, Jones GP (2001) Soft corals exert no 

direct effects on coral reef assemblages. 

Oecologia 127:560-571

Thomas AC, Weatherbee RA (2006) Satellitemeasured temporal variability of the 

Columbia River plume. Remote Sensing 

of Environment 100:167-178

Tissot BN, Yoklavich MM, Love MS, York K, Amend 

M (2006) Benthic invertebrates that 

form habitat on deep banks off southern 

California, with special reference to deep 

sea coral. Fishery Bulletin 104:167-181

Tissot BN, Wakefield WW, Puniwai NPF, Pirtle J, 

York K, Clemons JER (in prep) Abundance 

and distribution of structure-forming 

megafaunal invertebrates, including deep 

corals, on Heceta Bank, Oregon, 2000-

2002

Yoklavich MM, Love MS (2005) Christmas tree 

corals: A new species discovered off 

Southern California. Journal of Marine 

Education 21:27-30

Yoklavich MM, Greene HG, Cailliet GM, Sullivan 

DE, Lea RN, Love MS (2000) Habitat 

associations of deep-water rockfishes in 

a submarine canyon: an example of a 

natural refuge. Fishery Bulletin 98:625-

641

Wakefield WW, Whitmire CE, Clemons JER, 

Tissot BN (2005) Fish habitat studies:

combining high-resolution geological and 

biological data, In: Barnes PW, Thomas 

JP (eds) Benthic habitats and the effects 

of fishing: American Fisheries Society 

Symposium 41:119-138 

Wheatcroft RA, Sommerfield CK, Drake DE, 

Borgeld JC, Nittrouer CA (1997) Rapid and 

widespread dispersal of flood sediment on 

the northern California margin. Geology 

25:163-166

Williams GC (1995) Living genera of sea 

pens (Coelenterata: Octocorallia: 

Pennatulacea): illustrated key and 

synopses. Zoological Journal of the 

Linnean Society 113:93-140
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION Higher Taxon Coral Taxa San Diego Oregon Depth (m) Source Phylum Cnidaria Class Anthozoa Subclass Hexacorallia Order Scleractinia Family Fungiacyathidae Fungiacyathus marenzelleri Vaughan, 1906

x

x Cairns (1994)

Family Micrabaciidae Leptopenus discus Moseley, 1881

x Cairns (1994)

Family Rhizangiidae Astrangia haimei Verrill, 1866

x

x 1-53 Cairns (1994)

Family Oculinidae Oculina profunda Cairns, 1991

x 119 -578 Cairns (1994)

Madrepora oculata Linnaeus, 1758

x 84

Cairns (1994), Etnoyer and Morgan (2003)

Family Caryophylliidae Caryophyllia arnoldi Vaughan, 1900

x

x 183 - 505 Cairns (1994)

Labyrinthocyathus quaylei Durham, 1947

x

x 37 - 293 Cairns (1994)

Crispatotrochus foxi Durham and Barnard, 1952

x 82 Cairns (1994)

Paracyathus stearnsii Verrill, 1869

x

x Cairns (1994)

Paracyathus montereyensis Durham, 1947

x 75 - 146 Cairns (1994)

Coenocyathus bowersi Vaughan, 1906

x

x 9 - 302 Cairns (1994), NMFS

Nomlandia californica Durham and Barnard, 1952

x 82 Cairns (1994)

Desmophyllum dianthus (Esper, 1794)

x

x Cairns (1994)

Lophelia pertusa (Linnaeus, 1758)

x

x

Cairns (1994), Etnoyer and Morgan (2003), OCNMS, unpublished data

Family Flabellidae Javania californica Cairns, 1994

x 62 - 170 Cairns (1994)

Polymyces montereyensis (Durham, 1947)

x

x 69 - 212 Cairns (1994), NMFS

Family Dendrophylliidae Balanophyllia elegans Verrill, 1864

x

x Cairns (1994), Pirtle (2005)

Dendrophyllia oldroydae Oldroyd, 1924

x 99 - 366 Cairns (1994)

Appendix 3.1. Taxonomic list of known species of deep corals off the U.S. Pacific Coast, California to Washington. List includes both confirmed taxonomic records and some unconfirmed records of genera identified from surveys using bottom trawls and underwater vehicles. These unconfirmed records are denoted by the genus name followed by the word “species” (abbreviated as “sp”). Occurrence in two zoogeographic provinces (San Diego and Oregon) is noted. Known depth distributions are noted and originate from taxonomic records, published papers, and National Marine Fisheries Service (NMFS) bottom trawl surveys. Other abbreviations include the Olympic Coast (OCNMS) and Monterey Bay (MBNMS) National Marine Sanctuaries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

151PACIFIC COAST

Higher Taxon Coral Taxa San Diego Oregon Depth (m) Source

Order AntipathariaFamily Antipathidae Antipathes dendrochristos Opresko, 2005

x 90 - 360 Tissot et al. (2006)

Antipathes sp. 

x

x 82 - 1162

NMFS, Etnoyer and Morgan (2003), MBNMS, unpublished data

Family Cladopathidae Chrysopathes formosa Opresko, 2003

x Opresko (2003)

Chrysopathes speciosa Opresko, 2003

x 296 - 1400 Opresko (2003), NMFS

Trissopathes pseudotristicha Opresko, 2003

x

x

Opresko (2003), MBNMS, unpublished data

Family Schizopathidae Bathypathes sp. 

x

x 136 - 1243

NMFS, Etnoyer and Morgan (2003), MBNMS, unpublished data

Lillipathes sp. 

x 531 - 1243 NMFS, MBNMS, unpublished data

Umbellapathes sp. 

x MBNMS, unpublished data

Subclass OctocoralliaOrder GorgonaceaFamily Acanthogorgidae Acanthogorgia sp. 

x MBNMS, unpublished data

Calcigorgia spiculifera Broch, 1935

x 1127 - 1159 NMFS

Family Anthothelidae Anthothela pacifica (Kukenthal, 1913)

x

x Austin (1985)

Family Chrysogorgiidae Chrysogorgia sp. 

x MBNMS, unpublished data

Radiceps sp. 

x Austin (1985)

Family Coralliidae Corallium sp. 

x 1357 - 2447

MBNMS, unpublished data, OCNMS, unpublished data

Family Gorgoniidae Adelogorgia phyllosclera Bayer, 1958

x

x Montagne and Cadien (2001)

Eugorgia rubens Verrill, 1868

x Montagne and Cadien (2001)

Heterogorgia tortuosa Verrill, 1868

x Montagne and Cadien (2001)

Leptogorgia caryi Verrill, 1868

x

x 129 - 1200 Cairns et al. (2002), NMFS

Leptogorgia chilensis Verrill, 1868

x

x 86 - 710 Cairns et al. (2002), NMFS, OCNMS

Stenogorgia kofoidi Nutting, 1909

x Austin (1985)

Family Isididae Acanella sp. 

x MBNMS, unpublished data

Isidella sp. 

x

x 808 - 1165

NMFS, Etnoyer and Morgan (2003), MBNMS, unpublished data
PACIFIC COAST

152

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION Higher Taxon Coral Taxa San Diego Oregon Depth (m) Source Keratoisis flabellum Nutting, 1908

x Etnoyer and Morgan (2003)

Keratoisis philippinensis Wright and Studer, 1910

x Etnoyer and Morgan (2003)

Keratoisis sp. 

x

x 516 - 1707

NMFS, Etnoyer and Morgan (2003), MBNMS, unpublished data

Lepidisis sp. 

x

Etnoyer and Morgan (2003), MBNMS, unpublished data

Family Muriceidae Muricea californica Aurivillius, 1931

x Grigg (1970)

Muricea fruticosa Verrill, 1869

x Grigg (1970)

Thesea sp. A 

x

Ljubenkov (1986), Montagne and Cadien (2001)

Thesea sp. B 

x

Ljubenkov (1986), Montagne and Cadien (2001)

Family Paragorgiidae Paragorgia arborea (Linnaeus, 1758)

x 185 - 1743

Etnoyer and Morgan (2003), MBNMS, unpublished data, OCNMS, unpublished data

Paragorgia sp. 

x

x Etnoyer and Morgan (2003)

Family Plexauridae Swiftia spauldingi (Nutting, 1909)

x

x Austin (1985)

Swiftia beringi Nutting, 1912

x OCNMS, unpublished data

Swiftia kofoidi (Nutting, 1909)

x

x Austin (1985)

Swiftia pacifica (Nutting, 1912)

x 78 - 1186 NMFS, OCNMS, unpublished data

Swiftia simplex (Nutting, 1909)

x

x 62 - 1075 Austin (1985), NMFS

Swiftia torreyi (Nutting, 1909)

x

x Austin (1985)

Family Primnoidae Amphilaphis sp. 

x 114 - 114 NMFS

Callogorgia kinoshitae Kukenthal, 1913

x

x 127 - 464 NMFS, Etnoyer and Morgan (2003)

Calyptrophora sp. 

x Carey et al. (1990)

Narella sp. cf. bowersi Nutting, 1908

x Carey et al. (1990)

Parastenella doederleini Wright and Studer, 1889

x

x Etnoyer and Morgan (2003)

Parastenella sp. 

x Etnoyer and Morgan (2003)

Plumarella longispina Kinoshita, 1908

x Etnoyer and Morgan (2003)

Primnoa pacifica Kinoshita, 1907

x

x OCNMS, unpublished data
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION

153PACIFIC COAST

Higher Taxon Coral Taxa San Diego Oregon Depth (m) Source

Thouarella sp. 

x Etnoyer and Morgan (2003)

Order AlcyonaceaFamily Alcyoniidae Alcyonium rudyi Verseveldt and van Ofwegen, 1992

x Cairns et al. (2002)

Anthomastus ritteri Nutting, 1909

x

x 293 - 972 NMFS

Family Clavulariidae Cryptophyton goddarti Williams, 2000

x Cairns et al. (2002)

Clavularia sp. A 

x Austin (1985)

Clavularia sp. H 

x

Hochberg (1978), Montagne and Cadien (2001)

Telestula ambigua Nutting, 1909

x Austin (1985)

Telesto sp. 

x Montagne and Cadien (2001)

Family Nephtheidae Gersemia rubiformis (Ehrenberg, 1834)

x Austin (1985)

Order PennatulaceaSuborder SessilifloraeFamily Renillidae Renilla koellikeri Pfeffer, 1886

x Montagne and Cadien (2001)

Renilla reniformis (Pallas, 1766)

x intertidal Berntson et al. (2001)

Family Kophobelemnidae Kophobelemnon affine Studer, 1894

x Hochberg and Ljubenkov (1998)

Kophobelemnon biflorum Pasternak, 1960

x Austin (1985)

Kophobelemnon hispidum Nutting, 1912

x Austin (1985)

Family Anthoptilidae Anthoptilum grandiflorum (Verrill, 1879)

x

x 103 - 1543 Hochberg and Ljubenkov (1998), NMFS

Anthoptilum murrayi Kolliker, 1880

x

x 516 - 1083 NMFS

Family Funiculinidae Funiculina parkeri Kukenthal, 1913

x

x Hochberg and Ljubenkov (1998)

Family Protoptilidae Distichoptilum gracile Verrill, 1882

x Hochberg and Ljubenkov (1998)

Helicoptilum rigidum Nutting, 1912

x Austin (1985)

Family Stachyptilidae Stachyptilum superbum Studer, 1894

x

x Montagne and Cadien (2001)

Family Scleroptilidae Scleroptilum sp. 

x Austin (1985)

Family Umbellulidae Umbellula lindahli Kolliker, 1874

x Austin (1985)

Umbellula magniflora 

x Hochberg and Ljubenkov (1998)

Family Halipteridae Halipteris californica Moroff, 1902

x Hochberg and Ljubenkov (1998)
PACIFIC COAST

154

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC COAST REGION Higher Taxon Coral Taxa San Diego Oregon Depth (m) Source Halipteris sp. 

x

Suborder SubsellifloraeFamily Virgularidae Acanthoptilum album Nutting, 1909

x

x Hochberg and Ljubenkov (1998)

Acanthoptilum gracile (Gabb, 1863)

x Hochberg and Ljubenkov (1998)

Stylatula elongata (Gabb, 1862)

x

x Hochberg and Ljubenkov (1998)

Stylatula gracilis Verrill, 1864

x

x 64 - 1243 Hochberg and Ljubenkov (1998), NMFS

Stylatula sp. A 

x

Ljubenkov (1991), Montagne and Cadien (2001)

Virgularia agassizii Studer, 1894

x Montagne and Cadien (2001)

Virgularia californica Pfeffer, 1886

x Montagne and Cadien (2001)

Virgularia cystiferum (Nutting, 1909)

x

x Austin (1985)

Virgularia sp. cf glacialis Kolliker, 1870

x Austin (1985)

Family Pennatulidae Pennatula californica Kukenthal, 1913

x

x 465 - 991 Hochberg and Ljubenkov (1998), NMFS

Ptilosarcus gurneyi (Gray, 1860)

x

x 11 - 922 Hochberg and Ljubenkov (1998), NMFS

Ptilosarcus undulatus Verrill, 1865

x Cairns et al. (2002)

Class HydrozoaOrder AnthoathecataeSuborder FiliferaFamily Stylasteridae Errinopora pourtalesii (Dall, 1884)

x 49-183

Fisher (1938), Alberto Lindner, pers. comm.

Stylantheca petrograpta (Fisher, 1938)

x intertidal

Fisher (1938), Alberto Lindner, pers. comm.

Stylantheca porphyra Fisher, 1931

x intertidal

Fisher (1938), Alberto Lindner, pers. comm., Etnoyer and Morgan (2003)

Stylaster californicus (Verrill, 1866)

x

x 35-90

Fisher (1938), Alberto Lindner, pers. comm., Etnoyer and Morgan (2003)

Stylaster venustus (Verrill, 1868)

x 49-84

Fisher (1938), Alberto Lindner, pers. comm., Etnoyer and Morgan (2003)
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

155

STATE OF DEEP CORAL ECOSYSTEMS 

IN THE U.S. PACIFIC ISLANDS REGION: 

HAWAII AND THE U.S. PACIFIC TERRITORIES 

Frank A. Parrish1 and Amy R. Baco2

1

Pacific Islands Fisheries Science Center, 

NOAA

2570 Dole St. Honolulu, HI 96822

2

Woods Hole Oceanographic Institution

Biology Department

MS#33, 250 Redfield

Woods Hole, MA 02543

I. INTRODUCTION

The U.S. Pacific Islands Region consists of 

more than 50 oceanic islands, including two 

archipelagos (Hawaii and Mariana Islands), parts 

of four other archipelagos (Samoa, Line Islands, 

Phoenix Islands, and Marshall Islands), and 

numerous seamounts in proximity to each of these 

groups. These islands include the State of Hawaii, 

the Commonwealth of the Northern Mariana 

Islands (CNMI), and the territories of Guam and 

American Samoa, as well as nine sovereign 

Federal territories—Midway Atoll, Johnston Atoll, 

Kingman Reef, Palmyra Atoll, Jarvis Island, 

Howland Island, Baker Island, Rose Atoll, and 

Wake Island). This area also encompasses the 

Pacific Island States in free association with the 

United States (former U.S. trust territories also 

known as the Freely Associated States) including 

the Republic of Palau, the Federated States 

of Micronesia (Chuuk, Pohnpei, Kosrae, and 

Yap), and the Republic of the Marshall Islands. 

This region includes some of the most remote, 

unpopulated islands in the Pacific, as well as 

many densely populated islands, and it extends 

from the South Pacific (e.g., American Samoa; 

14o S latitude) to the North Pacific (Kure Atoll 28o 

N latitude) (Figure 4.1). The punctuated habitat of 

the Pacific Region distinguishes deepwater coral 

communities biogeographically and ecologically 

from other areas in the United States. Because 

of the isolated nature of these islands (especially 

Hawaii and the Northwestern Hawaiian Islands), 

they possess some of the highest levels of marine 

endemism recorded anywhere on earth.

While trace coral samples from anecdotal 

dredging and bycatch suggest a wide distribution 

of deep corals throughout the Pacific, the only 

detailed assessment of deep corals within the U.S. 

waters of the Pacific has been in the Hawaiian 

Archipelago. Antipatharians were first reported 

from Hawaiian waters more than 75 years ago 

(Verrill 1928). The earliest descriptions of deep 

octocorals in Hawaii are recorded by Dana (1846), 

with Nutting (1908) reporting 68 species. Other 

significant contributions to the species lists of this 

region include Muzik (1979) and Grigg and Bayer 

(1976) for octocorals, as well as Vaughan (1907) 

and Cairns (1984, 2006) for scleractinians. Wells 

(1954) provides data on the Marshall Islands. 

Pacific deep coral research has expanded greatly 

over the last four decades, primarily as a result 

of the establishment of commercial fisheries for 

black, pink, and gold coral off the main Hawaiian 

Islands, and subsequent development of fishery 

management plans for these resources by the 

State of Hawaii and the Western Pacific Fishery 

Management Council. Deep corals are harvested 

as raw material for the jewelry trade. The coral 

supports a portion of a $70 million Hawaii-based 

industry that employs roughly 650 people in its 

manufacturing facility and 50 retail stores (Carl 

Marsh—Maui Divers pers. comm.) 

Commercial beds of black coral were first 

discovered at a depth of 30-75 m off Lahaina, 

Maui in 1958. Some of the earliest ecological 

work on black corals was carried out in the 1960s 

in the channel waters off Maui using SCUBA 

(Grigg 1965). The Maui black coral bed has 

remained the focus of coral harvesters throughout 

the fishery’s history and has been periodically 

resurveyed over the last 40 years. These studies 

have provided the longest data sets available 

worldwide on the status and trends of black coral 

populations and the effects of the commercial 

fishery and other natural and anthropogenic 

stressors. In the mid 1960s, isolated patches 

of pink (Corallium spp.), gold (Gerardia sp.) 

and bamboo (Lepidisis sp., Acanella sp.) corals 
PACIFIC ISLANDS

156

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

were identified at 300–500 m depths north of 

Midway Island (Milwaukee Banks) and off Oahu 

(Makapuu Bed) (Grigg 1993). A long-term deep 

coral research program focused on precious 

corals began at the University of Hawaii in 1970. 

Many of the earliest surveys of precious coral beds 

used tangle net dredges and other nonselective 

gear. A key advancement in Hawaii’s deep coral 

research infrastructure was access to the twoperson submersible Deep Star 2 from General 

Dynamics. Aside from periodic research (Grigg 

1993) the sub was leased to commercially harvest 

coral for the fishery between 1974 and 1979. In 

1980 the submersible was renamed the Makalii

and became the centerpiece of the newly formed 

Hawaii Undersea Research Laboratory (HURL), 

an established node of NOAA’s Undersea 

Research Program (NURP). The facility has 

since expanded, replacing the Makalii with two, 

deep-diving 3-person submersibles (Pisces IV

and Pisces V) and a dedicated support vessel 

equipped with a multibeam bottom mapper and 

a remotely operated vehicle (RCV-150) (Chave 

and Malahoff 1998). This new infrastructure 

expanded the focus of coral research and 

increased participation by more researchers. 

This chapter provides a summary of what is 

known about deep corals within the Pacific 

Islands Region. In keeping with the intent of this 

national report, the chapter will mostly focus on 

corals deeper than 50 m. However, shallower 

black corals will be included. Most of the 

information available on black corals, precious 

corals, and other deep corals are from the 

Hawaiian Archipelago, where most of the surveys 

have been conducted. Studies have focused on 

the taxonomic and genetic composition of the 

region’s coral community, ecological relationships 

between corals and other organisms, and on the 

distribution and dynamics of deep corals. Much 

of this work is focused on the coral taxa that are 

targets for the commercial fishery. Also discussed 

are the measures that have been employed to 

Figure 4.1. Map of the Pacific Basin showing U.S. islands and their Exclusive Economic Zone that comprise 

the Pacific Islands Region for the National Marine Fisheries Service.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

157

protect deep coral ecosystems and to manage 

the commercial fishery. 

II. GEOLOGICAL SETTING

The U.S. Pacific Islands lack the shelf area that 

typically defines the deep-sea benthic habitats 

of the continental United States. Instead, the 

submerged slopes of volcanic pinnacles that rise 

steeply from abyssal depths of 4–7 km provide 

the hard substratum that deep corals colonize. 

The region has endured a long history of plate 

drift, subsidence, and sea level rise, and many 

of the volcanic islands have drowned creating 

numerous submerged banks and seamounts. A 

striking feature of the Pacific Basin is the linear 

nature of the island chains and seamounts. They 

are aligned in a north-northwesterly direction, 

a consequence of the northwesterly drift of the 

Pacific plate over stationary volcanic hotspots 

(Kennett 1982). The resulting islands and 

seamounts are progressively older in proportion 

to distance from a hotspot. For example, the 

island of Hawaii lies above the mantle plume and 

is the only island in the Hawaiian Archipelago that 

is volcanically active. To the northwest, volcanism 

on Oahu ceased about three million years ago; 

Kauai about five million years ago; and Midway 

Island about 27 million years ago (Grigg 1988a). 

This geologic process defines the Pacific plate 

and, as a result, the Pacific Islands region has 

some of the youngest (main Hawaiian Islands) 

and oldest (Line Islands) volcanic archipelagos 

in the world. 

III. OCEANOGRAPHIC SETTING

The Pacific is composed of two large gyres, 

the northern and southern central gyres. In the 

South Pacific, southeast trade winds drive the 

South Equatorial Current westerly between 15o

S and 3o N latitude. Within the South Equatorial 

Current is the Cromwell Current, or Pacific 

Equatorial Undercurrent. This current exists at 

depths of 70–200 m, and is approximately 450 

km wide and flows with velocities of up to 5 km h-1

for a distance of up to 13,000 km in the opposite 

direction of the South Equatorial Current (Tchernia 

1980; Thurman 1981). In the North Pacific, the 

North Equatorial Current flows westward at 1 km 

h-1 between 8o and 20o N latitude. The Equatorial 

Counter Current is located between the North 

Equatorial Current and the South Equatorial 

Current and travels eastward at slightly more 

than 2 km h-1 (Thurman 1981). The boundaries 

of these water masses overlap and contribute 

to long-distance dispersal of pelagic larvae, 

particularly in the western Pacific. The northern 

Hawaiian seamounts fall in the northern portions 

of the north gyre; Hawaii, Wake and Johnston are 

in the center of the North Pacific gyre; Kingman 

and Palmyra are in the equatorial/eastern Pacific; 

Jarvis, Howland, and Baker are in the equatorial 

system; American Samoa is in the equatorial 

portion of the southern gyre; and the Mariana 

Islands are affected by the north central gyre, 

the equatorial systems, and the Kuroshio current 

from Asia. 

While deep water masses originate from surface 

currents, no deep water masses form in the 

Pacific Basin. Deep water migrates in from 

the Atlantic via the southern hemisphere with a 

uniform temperature and salinity below about 

2000 m (Knauss 1996). The deep water flows 

northward at depths below 2500 m and southward 

above 2500 m. Seamounts, pinnacles, and other 

structures obstruct current flow and can generate 

eddies of varying intensity, depending on the 

current velocity, depth or height of the seamount, 

slope of the side, and strength of the seawater 

stratification. Both cold and warm water eddies 

are formed as a result of a seamount obstructing 

current flow in the deep ocean. Typically, 

anticyclonic (cold water) eddies are formed above 

the seamount and remain tightly associated 

with the top of the seamount, while the cyclonic 

(warm water) eddy is formed downstream 

behind the seamount (Kamenkovich et al. 

1986). Deep corals are thought to benefit from 

the flow acceleration, larval retention, and high 

nutrient waters from deep upwelling caused by 

the presence of the seamount and the generated 

eddies (Genin et al. 1986; Mullineaux and Mills 

1997; Coutis and Middleton 2002). 

Oxygen in the deep water of the Pacific has been 

depleted to very low levels as a result of the length 

of time it takes for water to move into and across 

the Pacific Basin. Having aged thousands of 

years, oxygen averages 0.5–4.5 ml l-1 versus the 

significantly higher Atlantic average of 3.0–6.5 

ml l-1 (Thurman 1981). The effect of low oxygen 

levels on deep corals is poorly documented; 

however, Wishner et al. (1990) attributed patterns 

in the abundance and distribution of sponges, 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

sea pens, and other benthic organisms to depthspecific patterns in oxygen levels.

IV. STRUCTURE AND HABITAT-FORMING 

DEEP CORALS

Most of the major deep coral groups are known 

to exist in the U.S. Pacific region. However, 

most species have been identified only around 

the Hawaiian Archipelago largely because deep 

waters around other U.S Pacific Islands have not 

yet been explored. Published records of deep 

corals from the Hawaiian Archipelago include more 

than 137 species of gorgonian octocorals and 63 

species of azooxanthellate scleractinians, with 

21% of the scleractinians thought to be endemic 

to Hawaii (Cairns 2006, See chapter appendix). 

A 2003 cruise in the Northwestern Hawaiian 

Islands (A. Baco, unpublished data) identified 

eight new species of octocorals, two new genera 

and several new species of antipatharians, three 

new stylasterid species (Cairns 2005), and 1 

new zoanthid species. In addition, a 2004 main 

Hawaiian Islands cruise collected at least three 

new species of octocorals, two new species of 

Taxa

ReefBuilding Abundance

Maximum 

Colony 

Size Morphology

Associations 

with Other 

StructureForming 

Invertebrates

Colony 

Spatial 

Dispersion

Overall 

Rating of 

Structural 

Importance

Enallopsammia 

rostrata No Medium Medium Branching Many Clumped Medium

Other 

scleractinians No Low Small

NonBranching Many Solitary Low

Gerardia sp. No High Large Branching Many Clumped High

Shallower 

antipatharians No Low Large Branching Few Clumped High

Other 

octocorals and 

antipatharians in 

deeper water No High Med Branching Many Clumped High

Other 

octocorals and 

antipatharians 

in precious coral 

beds No Medium

Medium 

-Low Branching Many Clumped High

Corallium 

secundum No High Medium Branching Many Clumped Medium

Corallium 

laauense No High Medium Branching Many Clumped Medium

Isidids in deeper 

water No High Med Branching Many Clumped Medium

Table 4.1 Structure-forming attributes of deep corals in Hawaii.

Table Key

Attribute Measure

Reef-Building Yes/No

Relative Abundance Low/ Medium/ High

Size (width or height) Small (<30cm)/ Medium (30cm-1m)/ Large (>1m)

Morphology Branching/ Non-branching

Associations None/ Few (1-2)/ Many (>2)

Spatial Dispersion Solitary/ Clumped

Overall Rating Low/ Medium/ High
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

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159

antipatharians, and provided range extensions 

for several genera and species of corals that 

were not previously known from Hawaii. Thus, 

although an extensive species list exists for the 

Hawaiian Archipelago, the high rate of discovery 

of new species and new records implies the 

Archipelago is also largely undersampled.

In general, the deep corals in the Pacific Islands 

do not form the extensive reef structures observed 

in the Atlantic and South Pacific. Instead, corals 

grow attached directly to the exposed fossil 

carbonate, basalt or manganese substratum. 

Octocorals and antipatharians have been found 

to grow in high densities at numerous sites, 

particularly on summit areas of seamounts or 

other topographic highs, where they often form 

extensive coral gardens or “beds” with abundant 

associated invertebrates. The coral taxa listed in 

this chapter are those that present a conspicuous 

relief profile that could serve as a source of 

habitat (Table 4.1). At death, these taxa decay 

from physical and bioerosion forces until they 

detach from the substratum and are swept away. 

Cemented reefs from accumulated dead material 

have not been observed. 

a. Stony corals (Class Anthozoa, Order

 Scleractinia)

Enallopsammia rostrata is an arborescent 

scleractinian coral in the Family Dendrophyllidae. 

The full depth range for this species is listed as 

229–2165 m in Cairns (1984), but it has been 

observed in Hawaii primarily at depths of 500–

600 m. In some areas it forms bushy colonies, 

with dead coral accumulating near the base 

of the colony much like that observed among 

Lophelia reefs in the Atlantic. Further exploration 

and characterization of this species is needed to 

determine its abundance throughout the region 

and its potential role in forming habitat.

Madrepora kauaiensis and M. oculata also 

occur in Hawaii and have the potential to form 

reef structures, however, little is known of their 

abundance or distribution in the Archipelago. 

Besides these examples, scleractinians that 

have been observed are primarily solitary cup 

corals. They can occur in abundance, e.g., on 

Cross Seamount (A. Baco pers. obs), but many 

species are small and not observable with a 

submersible, preventing a true determination of 

their distribution. A complete species list (to date) 

for Hawaii can be found in Cairns (2006) and is 

also included in the Appendix to this chapter. 

b. Black corals (Class Anthozoa, Order

 Antipatharia)

Fourteen genera of black corals are reported from 

the Hawaii-Pacific region with species found in 

both shallow and deep habitats. The shallowest 

genera (Cirripathes spp. and Antipathes spp.) 

prefer shaded or low light areas and can occur 

underneath ledges and in caves in shallow water 

(e.g., Cirrhipathes anguina can occur in 4 m depth) 

where surge is minimal, or in the open on steep 

walls at deeper depths. Antipathes spp. appears 

to settle predominantly in depressions, cracks or 

other rugged features along steep ledges, with 

few colonies found on smooth basaltic substratum 

(Grigg 1965). Shallower antipatharians in 

Hawaii also appear to prefer substrates that are 

encrusted with calcium carbonate from coralline 

algae, bryozoans, and corals. The highest 

densities are found on hard sloping substratum, in 

areas with 0.5–2 knot currents (Grigg 1965). The 

best studied black corals are the commercially 

harvested species Antipathes dichotoma and

Antipathes grandis. Recent taxonomic work 

(D. Opresko pers. comm.) on the Hawaiian A. 

dichotoma suggests it is a new species and is 

currently being referred to as Antipathes cf. 

curvata. The A. cf. curvata and A. grandis exhibit 

similar growth rates (6.42 cm yr-1 and 6.12 cm yr-1, 

respectively) and reach reproductive maturity at 

ages 12–13. Fertilization takes place externally 

in the water column, and light and temperature 

influence larval settlement patterns. The larvae 

of A. cf. curvata and A. grandis are negatively 

phototactic, and the lower depth limit coincides 

with the top of the thermocline (~ 100 m) in the 

main Hawaiian Islands (Grigg 1993). 

Much less is known about deeper genera of 

black corals. They have similar morphologies to 

the shallower corals, including colonies shaped 

as whip-like filaments (Stichopathes spp.) and 

as branching, sometimes “feathery” colonies 

(Myriopathes, Bathypathes spp., Stauropathes 

and Leiopathes). However, the life history of 

these deeper genera is likely to be much different 

than their shallower relatives. Radiometric dating 

of A. cf. cruvata and Leiopathes indicates the 

deeper Leiopathes genera grow 10 to 70 times 

slower than the shallower A. cf. curvata (Roark 

et al. 2006). 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

c. Gold coral (Class Anthozoa, Order 

 Zoanthidea)

No species of zoanthid has yet been described 

from deep water in Hawaii although taxonomically, 

at least six species have been observed 

and,ollected (Baco, unpublished data). The gold 

coral, Gerardia sp., is probably the most common 

and certainly the largest of these species. It has 

an arborescent morphology similar to gorgonians, 

and colonies have been observed as tall as 2–3 

m in height (Figure 4.2 B). Gerardia sp. is widely 

distributed throughout the Hawaiian Archipelago 

and into the Emperor Seamount Chain at depths 

of 350–600 m.

Zoanthids in Hawaii have been observed to 

colonize other living coral species as well as 

hard bottoms. In the case of Gerardia sp., the 

zoanthids may eventually replace the host colony 

completely. It is not known if Gerardia sp. can 

outcompete the living tissue of the host or if it 

opportunistically colonizes and spreads after a 

portion of the host coral has been decorticated 

by predatory urchins or some other cause. The 

life span of Gerardia sp. is uncertain. Counts of 

growth bands assumed to be annual in periodicity 

have provided an estimated lifespan of around 40 

years (Grigg 2002). Recent radiometric work on 

the Hawaiian species has estimated the life span 

of gold coral samples between 450 and 2700 

years (Roark et al. 2006), which is consistent 

with findings from radiometric aging on Gerardia

sp. in the Atlantic (Druffel et al. 1995). 

Gold coral was one of the few corals seen during 

the 2005 Line Island surveys and it was present 

at Jarvis, Palymra, and Kingman. However, the 

colonies were sparse, with no patches large 

enough to be called a “bed.” All colonies were 

infested with other unidentified zoanthids. Surveys 

at the base of the cliffs below the spot where the 

individual gold colonies were attached found no 

accumulation of fallen colonies, suggesting gold 

coral has always been in low abundance in the 

region (Frank Parrish pers. obs.). 

d. Gorgonians (Class Anthozoa, Order 

 Gorgonacea) 

Gorgonian octocorals are by far the most 

abundant and diverse corals in the Hawaiian 

Archipelago. Two species, Corallium laauense 

(red coral; formely identified as Corallium regale) 

and Corallium secundum (pink coral) are known 

to occur at depths of 350–600 m on islands and 

seamounts throughout the Hawaiian Archipelago 

(Grigg 1974, 1993; Baco, unpublished data) 

and into the Emperor Seamount Chain (Bayer 

1956). Growing to more that 30 cm in height 

the Corallium spp. occur in a variety of red/pink 

color shades, and the height and shape of the 

colony’s “fan” can vary considerably (Figure 

4.2 A). They are often found in large beds and 

usually support a high diversity of invertebrates 

with an abundance of other octocorals, zoanthids, 

and sometimes scleractinians co-occuring in 

the beds. C. secundum and C. laauense are 

gonochoristic (Grigg 1993; Waller and Baco in 

press) and are estimated to reach reproductive 

Figure 4.2. Photos of the two primary Genera that comprise the Hawaiian precious coral fishery 

A—Corallium sp. and B—Gerardia sp. Photo credit F. Parrish, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

161

maturity at 12–13 years (Grigg 1993). These 

species are relatively long lived, with some of the 

oldest colonies observed within Makapuu Bed 

about 0.7 m in height and approximately 80 years 

old (Grigg 1988b, Roark 2006). Populations of 

C. secundum appear to be recruitment limited, 

although in favorable environments (e.g., 

Makapuu Bed) populations are relatively stable, 

suggesting that recruitment and mortality are in a 

steady state (Grigg 1993). 

More than 130 other species of octocorals are 

known from the Archipelago and they represent 

a diverse array of families and genera. Most 

abundant are the Families Coralliidae, Isididae, 

Primnoidae, Plexauridae, Chrysogorgiidae and 

Acanthogorgiidae. To our knowledge, besides 

the species discussed above, there isn’t any 

information on the biology and ecology of these 

groups in this region. Further discussion of their 

depth distributions are in the following sections 

and a complete species list (to date) is included 

in the Appendix to this chapter.

e. True soft corals (Class Anthozoa, Order 

 Alcyonacea)

The Alcyonacea are represented in this region by 

only 12 species in three families. Of these, the 

genus Anthomastus is the most widely distributed. 

It is often observed in precious coral beds in 

patches with large number of small individuals 

surrounding a larger individual (A. Baco pers. 

obs.). The bright purple Clavularia grandiflora

has been observed growing on Gerardia at a 

number of sites throughout the Archipelago (A. 

Baco pers. obs.) 

Like the gorgonians, little else is known about 

the biology and ecology of these species in 

this region. A complete species list (to date) is 

included in the Appendix to this chapter. 

f. Pennatulaceans (Class Anthozoa, Order 

 Pennatulacea)

Pennatulaceans seen in Hawaii tend to be more 

abundant in areas high in sediment, although 

Kophobelemnon sp. has occasionally been 

observed on adjacent hard bottoms near the 

Cross Seamount deep coral bed. Near the 

Keahole deep coral bed on the island of Hawaii, 

a single species (as yet unidentified) occurs in 

high abundance in patches of sediment at depths 

of about 400 m (A. Baco pers. obs.). Again, little 

else is known about the biology and ecology of 

these species in this region. A complete species 

list (to date) is included in the Appendix to this 

chapter. 

g. Stylasterids (Class Hydrozoa,

 Order Anthoathecatae)

Four species of stylaserids are present in 

Hawaii, but they are distributed very patchily 

throughout the Archipelago. An extreme example 

is Disticophora anceps. It has a very wide depth 

range but has only been found on the northwest 

slope of Laysan Island in densities of several 

colonies per square meter in some areas (Cairns 

2005; A. Baco unpublished data). Again, little 

else is known about the biology and ecology of 

this group in this region.

Hawaiian stylasterids are discussed in Cairns 

(2005) and a complete species list (to date) is 

included in the Appendix to this chapter.

V. SPATIAL DISTRIBUTION OF CORAL 

SPECIES AND HABITAT

General distribution

Our knowledge of the spatial distribution of deep 

corals in the U.S. Insular Pacific is limited to 

Hawaii. Even in Hawaii, very little of the deep 

sea has been explored and every research 

expedition is yielding large numbers of new 

species. Until 2003, the majority of studies in 

Hawaii came from sparse trawl data or had 

concentrated on the harvested black, gold, and 

pink corals. Often referred to as “precious corals,” 

these are the primary deep coral taxa harvested 

for the jewelry trade. Most of these are found 

between depths of 300 and 500 m and have 

been collected by dredge or submersible. Often, 

black coral (Antipathes spp.) is distinguished 

from the rest of the precious corals because the 

Antipathes taxa that are used for jewelry occur 

much shallower (<100 m) and are harvested 

by scuba divers. Beside precious corals, many 

other taxa of deep corals have not been studied 

because they were not one of the management 

unit species of the precious coral fishery. In 2003 

and 2004, the Pisces submersibles were used 

for the first studies of non-precious corals, thus 

extending the exploration of corals well below 

previously surveyed depths. In 2005, the first 

surveys conducted outside of Hawaii using the 

Pisces submersibles were at Rose Atoll and the 

U.S Line Islands (Jarvis, Palmyra, and Kingman). 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

The distribution of deep corals in the rest of the 

U.S. Pacific is unknown.

Depth clearly influences the distribution of 

different coral taxa and certainly there is 

patchiness associated with the presence of 

premium substrate and environmental conditions 

(flow, particulate load, etc.). The environmental 

suitability for colonization and growth is likely to 

differ among coral taxa. For example, Corallium 

secundum appears to grow in large numbers 

in areas of high flow over carbonate pavement; 

Corallium laauense grows in an intermediate 

relief of outcrops; and Gerardia sp. grows in high 

relief areas on pinnacles, walls, and cliffs (Parrish 

in press). These habitat differences may reflect 

preferred flow regimes for the different corals 

(e.g., laminar flow for C. secundum, alternating 

flow for Gerardia sp.). 

Black coral beds

Black coral beds are found off the main Hawaiian 

Islands at depths of about 30–110 m. The largest 

bed covers an estimated area of 1.7 km2 and lies 

in the middle of the Auau Channel, between Maui 

and Lanai, encrusting a drowned land bridge 

between the two islands at depths of 30-90 m 

(Grigg et al. 2002). A smaller black coral bed 

(0.4 km2) is located off Kauai and another at the 

southern end of the island of Hawaii (Figure 4.3). 

The dominant species found in these locations 

are Antipathes cf. curvata (95% of the population) 

followed by Antipathes grandis. Grigg (1976) 

estimated a total standing crop of A. cf. curvata

for the Auau Channel area, between 40 and 70 

m, to be 166,000 kg or 84,000 colonies, while the 

bed at Kauai contained 40,000 kg. Myriopathes 

ulex is found in deeper locations (110–565 m) 

off the main Hawaiian Islands, along with other 

species of antipatharians absent from shallower 

depths (Devaney and Eldridge 1977) (see 

chapter appendix). Little commercially important 

black coral has been found in the Northwestern 

Hawaiian Islands (Grigg 1974), perhaps due to the 

shoaling of the thermocline towards the northwest 

end of the chain. Other species of black corals 

Figure 4.3. Topographic map of the main Hawaiian Islands with the three known black coral beds marked. 

Inset shows a diver conducting a coral survey. Map creidt: F. Parrish, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

163

occur in the Northwestern Hawaiian Islands and 

include taxa such as Cirripathes, Stichopathes, 

Stauropathes, Bathypathes, Myriopathes ulex, 

Trissopathes, Umbellopathes, Dendropathes, 

and Leiopathes. 

Between Black and Precious Coral Beds

The depth zone between the black coral beds 

and the precious coral beds has had less study. 

Corals have been observed in this zone; for 

example, an abundance of octocorals occur at 

the Makapuu coral bed on the island of Oahu, 

shallower than the precious corals, but they have 

not been sufficiently sampled to comment on 

diversity or species composition. 

Precious coral beds

Probably the most abundant of Hawaii’s known 

deep corals are the precious corals, including 

octocorals Corallium laauense (red coral) and 

Corallium secundum (pink coral), and the 

zoanthid Gerardia sp. (gold coral). These species 

are known to occur in significant abundance in 

at least 16 locations in the Hawaiian Archipelago 

at depths of 350–600 m (Grigg 1974, 1993; 

Baco, unpublished data) and into the Emperor 

Seamount Chain (Bayer 1956). Within a given 

coral bed, the two primary genera (Corallium

and Gerardia) are usually found, but the ratio 

of abundance can vary greatly (Parrish in 

press)(Figure 4.4). It is difficult to estimate the 

size of coral beds, so only relative differences in 

bed size (based on impressions of coral density 

and the area covered by the submersible track) 

were shown in Figure 4.4 to determine the size of 

pie diagrams. Most precious coral sites also have 

a number of other noncommercial taxa; these 

include various octocorals (e.g., Callogorgia, 

Paracalyptrophora, Acanthogorgia, Lepidisis, 

Keratoisis, Isidella, Kereoides, Paragorgia, and 

various paramuriceids) and antipatharians (e.g. 

Leiopathes, Trissopathes, and Bathypathes) (A. 

Baco, unpublished data). 

Of the known coral beds, the Makapuu coral bed 

is the best studied and most diverse. It is located 

between 375 and 450 m depth in the channel 

between the islands of Oahu and Molokai. The 

bed comprises an area of about 3.6 km2, with 

the most abundant coral C. secundum, at a 

mean density of 0.22 colonies per square meter 

between 365 and 400 m (Grigg 1988b). Other 

corals found at Makapuu include bamboo coral 

(Lepidis olapa, 0.041 colonies m-2; Acanella spp., 

0.001 colonies m-2), gold coral (Gerardia sp., 

0.0005 m-2), as well as three genera of gorgonians 

Figure 4.4. Topographic map of the Northwestern Hawaiian Islands with coral survey dive sites of the submersible Pisces V (inset). Pie charts represent the relative amount of coral among sites and the within-site 

fraction of the two primary precious coral genera. 3D map and Pisces photo credit: Hawaii Undersea Research 

Laboratory.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

(Narella sp., Psuedothesea sp and Callorgorgia 

gilberti), a sea pen (Stylatula sp.), and black coral 

(Leiopathes n. sp.). Many taxa of no commercial 

interest are also present in or adjacent to the bed 

including Enallopsammia rostrata, Thouarella 

hilgendorfi, acanthogorgiids, Paragorgia sp., 

Paramuriceidae, Trissopathes pseudotristicha

(antipatharian) and a number of undescribed 

octocorals. 

Beyond Precious Coral Beds

Explorations have been conducted at only a 

few sites below precious coral depths; Pioneer 

Ridge, the small seamount southeast of Laysan 

Island, an unnamed seamount east of Necker 

Island, Cross Seamount, and Keahole Point. At 

these deeper depths in high current areas such 

as ridges and pinnacles, a fair amount of overlap 

appears to occur in species composition of both 

corals and sponges between sites. Although 

the number of observations is very limited, there 

appears to be a transition in species below about 

600 m, from Corallium- and Gerardia-dominated 

communities, to a different suite of species. Many 

species of chrysogorgiids, primnoids, isidids, 

coralliids, and antipatharians begin to appear 

around this depth (A. Baco, unpublished data). 

Among the more common octocoral genera 

observed are: Chrysogorgia, Metallogorgia, 

Iridigorgia, Narella, Calyptrophora, Candidella, 

Keratoisis, Isidella, Acanella, Corallium, and 

Paragorgia, as well as the antipatharian genus 

Bathypathes. The depth distribution of many, 

but not all, of these species appears to continue 

below 1800 m (Baco, unpublished data). 

 

V. SPECIES ASSOCIATIONS WITH DEEP

CORAL COMMUNITIES

The harvesting of deep corals has prompted 

a number of studies focused on the species 

associations with deep coral communities. 

Nationally, most effects to coral-associated 

species come from trawling and dredging, 

which are banned in U.S. Pacific waters. The 

harvesting of precious corals is allowed using 

selective methodologies such as hand collection 

by scuba divers or using the manipulator of a 

submersible. Take of deep corals as a fishery 

target is a direct effect to the bottom habitat with 

uncertain ecological consequences. Studies have 

been conducted to address the NOAA mandate 

of essential fish habitat, protected species, and 

ecosystem concerns. These have historically 

been focused on fish and only recently have been 

expanded to include invertebrates. 

Commercial fishery species

With the exception of the fishery that harvests 

precious corals, there is little evidence of a direct 

association between precious corals and other 

fishery targets. However, these evaluations 

have been limited to comparing the overlap in 

depth ranges and making infrequent underwater 

observations. Even less is known about the deep 

sea corals not targeted in 

the precious coral fishery, 

or any indirect ecological 

effects that any of these 

corals may contribute to 

commercial fishery stocks. 

Some of the shallow 

coral reef fish targeted by 

recreational fishers and the 

aquarium trade range into 

depths where black corals 

(antipatharians) can be 

found (30–100 m) (Moffitt 

et al. 1989; Parrish and 

Boland 2004; Boland and 

Parrish 2005). One of the 

commercially sought bottom 

fish Aprion virescens (grey 

snapper), also lives in this 

depth range but most of 

the commercial bottom 

Figure 4.5. Two species of black coral trees Antipathes grandis (left) and 

Antipathes. cf. curvata (right). Photo credit F. Parrish, NOAA Fisheries.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

165

fish reside at depths below antipatharians and 

above the precious corals (<300 m) (Uchida 

and Tagami 1984). The shallowest members of 

this group, such as Pristipomoides filamentosus 

(pink snapper), Pristipomoides zonatus (obliquebanded snapper), and Epinephelus quernus

(Hawaiian grouper), have been seen in the 

vicinity of deeper black coral trees (Moffitt et al. 

1989). Similarly, the deeper members including 

Etelis carbunculus (ruby snapper) and Etelis 

coruscans (flame snapper) have been seen 

among the shallower precious corals (Kelley et 

al. 2006). The groundfish, Pseudopentaceros 

wheeleri (armorhead) and Beryx sp. (alfonsino) 

occur throughout this depth range (250–350) 

but are more common on the seamounts at the 

northern end of the Hawaiian Archipelago (Uchida 

and Tagami 1984). There is no information on 

the degree of overlap of these fish with deep sea 

corals. Heterocarpus sp. (deep-water shrimp) 

has been seen among the precious corals but at 

densities consistent with other bottom relief types. 

Heterocarpus sp. is the focus of an intermittent 

main Hawaiian Island trap fishery that targets 

depths of 500 to 900 m (Moffitt and Parrish 1992) 

overlapping the lower limit of precious coral 

depths, but in the depth range of many other 

deep coral species. 

Noncommercial species 

Fish

Studies of fish associations with deep corals have 

focused almost exclusively on the inventory of fish 

taxa and appraisal of the obligate or facultative 

roles corals play in fish assemblages. The fish 

community of the Auau Channel black coral bed 

was recently surveyed (Boland and Parrish 2005) 

and 95% of the fish found in and around the 

black corals were known to commonly occur on 

shallower reefs where black corals do not grow. 

Oxycirrhites typus (the longnose hawkfish), was 

found exclusively within the black coral trees. 

Behavioral data indicated that although most of 

the reef fish routinely passed through the coral 

Figure 4.6. Examples of invertebrates found associated with deep corals (A-basket star, B-ophiuroid, 

C-asteroid. Photo credit A.Baco, WHOI.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

branches, only four species reliably used the 

corals for cover when evading a threat (Figure 

4.5). Although there is little or no known obligate 

relationship between fish and black coral, the 

coral colonies contribute to the fish community 

by enhancing the vertical aspects of the deep 

reef and perhaps improving the corridors for fish 

movement. 

Most of the fish of the deep slope and subphotic 

depths are noncommercial species (Chave and 

Mundy 1994). The surveys of fish communities 

at deeper subphotic depths indicated few fish 

associations with precious corals (Parrish 2006). 

Many of the 42 fish taxa observed were seen 

to use Gerardia and Corallium spp. as shelter 

interchangeably with abiotic relief sources. 

Species richness of fish was not observed to differ 

between areas with corals and those without. 

Most fish taxa were observed in low numbers 

with only a couple of dominant species. Areas 

with corals often supported greater fish density, 

but statistical evaluations suggested that this 

was based on co-occurrence of fish and coral in 

areas of relief and high flow and not based on a 

dependency of fish on corals. Also, differences 

were not seen in the mean size of fish in or outside 

of the coral beds. Of the three commercial coral 

species, fish oriented mostly around Gerardia sp. 

probably because it is significantly taller than the 

two Corallium species. Classifying the fish into 

functional groups revealed “benthic hoverers” 

as the segment of the fish community that most 

frequently used Gerardia sp. as shelter. 

Although use of corals by the fish appeared 

incidental, it is important to consider these studies 

were conducted in summer, during the day and 

focused exclusively on adult fish. It is unknown 

if the corals play some seasonal or diurnal role in 

the fish ecology or if juvenile stages rely on the 

coral colonies.

Invertebrates 

There is a wide array of invertebrate species 

associated with deep corals. Black corals are 

Figure 4.7. The 

purple octocoral 

Clavularia grandiflora growing 

on a Gerardia sp. 

colony.Photo credit 

A. Baco, WHOI.
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known to be colonized by oysters, bryozoans, 

and shrimp (Hoover 1998). The most common 

invertebrates associated with the deeper precious 

corals include zoanthids, anemones, galathaeoid 

crabs, sponges, ophiuroids, and basket stars 

(Figure 4.6). Gerardia sp. is perhaps the best 

known coral that overgrows the skeletons of 

other coral species. Examination of Gerardia sp. 

skeletons and many submersible observations 

suggests it has an obligate need to start its 

colony over the skeleton of other corals. Bamboo 

corals seem to be the most frequent target, 

particularly Isidella trichotoma, but Gerardia has 

been observed growing on quite a number of 

species. Several other species of unidentified 

zoanthids also appear to prefer to grow on other 

corals, including Gerardia sp. At least one of 

these zoanthids was observed growing on basalt 

substrate as well as encrusting other corals (A. 

Baco pers. obs.). Octocorals also can grow 

on the skeletons of other coral species. The 

bright purple Clavularia grandiflora has been 

observed growing on Gerardia at a number of 

sites throughout the Archipelago (Figure 4.7). 

In all of these cases, it is not clear whether the 

overgrowing corals are actually killing or injuring 

the coral whose skeleton they are growing on, or 

if they have simply colonized a dead area of the 

skeleton. 

Many coral colonies have a number of commensal 

invertebrates such as galathaeoid crabs, 

barnacles, sponges, anemones, polychaetes, 

crinoids, ophiuroids, and basket stars (Chave and 

Malahoff 1998; A. Baco and Shirley, unpublished 

data). Casual observations suggest some of these 

commensals may be specific to their host while 

others are more generalized. The relationships 

between the coral and the commensal, e.g., 

symbiotic, parasitic, obligate, facultative, etc., 

have not been determined in most cases and 

many of the commensal species have yet to be 

identified. One of the more notable commensal 

relationships is the general association of 

polychaetes with species in the genus Corallium. 

Each Corallium species appears to have its own 

species of polynoid polychaete. In Corallium 

secundum and Corallium laauense, these 

polychaetes can reach fairly high densities. The 

polychaetes live in tunnels under the coral soft 

tissue with the skeleton often growing over the 

polychaete tunnels. This relationship has also 

been observed in other corals such as Candidella 

helminthophora. 

A more generalized commensal is the unbranched 

basket star, Asteroschema. Asteroschema sp. 

has been observed in a number of different species 

of octocorals at a number of sites, but not on the 

surrounding substratum. Dead coral skeletons 

also appear to provide good recruitment habitat 

for many invertebrate species. Many types of 

sessile fauna have been observed as well as 

several types of young corals. In particular, young 

colonies of the red coral, Corallium laauense

have been observed growing on dead skeletons 

of Gerardia sp. 

Besides the galathaeoid crabs that inhabit the 

branches of coral colonies, a number of larger 

crabs are routinely encountered patrolling the 

bottom around deep corals (e.g., crabs in the 

families Homolidae, Parapaguridae). 

Predation on deep corals by resident invertebrates 

also occurs. Seastars feed on coral colonies 

by everting their gut, leaving behind patches 

of bare coral skeleton. Cidarid urchins are 

also known to feed on deep corals and these 

urchins have been observed on deep corals 

in Hawaii. However, an absence of bare coral 

skeleton around the urchin’s location suggests it 

is unlikely they are consuming the coral tissues 

(A. Baco, unpublished data). A single species of 

orange crinoid was observed in 2004 in very high 

densities at the Makapuu coral bed (A. Baco, 

unpublished data). Grigg 2002 comments on an 

abundant crinoid observed during night dives at 

this site. The crinoids now cover many octocoral 

colonies and anything else that sticks up more 

than a few millimeters off the bottom (A. Baco, 

unpublished data), suggesting their abundance 

has increased over Grigg’s observations. Their 

origin and the reason for their recent increase in 

population density, as well as their potential for 

competing with corals for food, are unknown. 

Monk seals

In the lower Northwestern Hawaiian Islands, 

the endangered Hawaiian monk seal has been 

documented to routinely visit deep corals as 

part of its foraging activities. However, there 

have been no reported interactions between 

monk seals and the precious coral harvesting 

in the main Hawaiian Islands. Telemetry and 

scatological analysis indicate seals prey on 

bottom-dwelling fish (Goodman-Lowe 1998; 

Parrish and Abernathy 2006; Longnecker et 

al. 2006). Video cameras fitted to seals in the 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

Table 4.2 Potential effects of fishing gears on deep coral habitat in Hawaii. Ratings detailed in table key. Tuna/

Swordfish includes: Albacore, Bigeye tuna, Yellowfin Tuna, Skipjack Tuna, Swordfish, Striped Marlin, Pacific 

Blue Marlin, Black Marlin Sailfish, Shortbill spearfish, Wahoo, Dolphinfish, Opah, Pomfret, sharks. Deepwater 

shrimp include: two species of Heterocarpus, Snappers and Groupers include: Pink snapper, Flower snapper, 

Squirrelfish snapper, Hawaiian Grouper, Ruby-colored snapper, Blue-green snapperfish.

Gear Type

Current 

Fishery 

Use in 

Region

Potential 

Severity 

of Impact

Potential 

Extent of 

Impact 

from 

Fishing 

Gear

Current 

Geographic 

Extent 

of Use in 

Region

Overlap 

of use 

with coral 

habitat

Overall 

Rating 

of Gear 

Impact

Bottom Trawl N/A High High N/A N/A N/A

Mid-water Trawl N/A Low Low N/A N/A N/A

Dredge N/A High Low N/A N/A N/A

Bottom-set 

Longline N/A Med Low Low N/A N/A

Bottom-set 

Gillnet N/A Med Med N/A N/A N/A

Pelagic longline Tuna/ 

swordfish Low Med Med Low Low

Traps Deepwater 

Shrimp Med Med Low Low Low

Hook and line Snappers/ 

groupers

Low Low High Low Low

Northwestern Hawaiian Islands have recorded 

seals commuting to beds of Cirripathes sp. 

whip corals (100 m) where they feed on eels. 

Satellite tags attached to seals indicated certain 

seals spent weeks of their foraging focused at 

specific subphotic locations where surveys with 

submarines have revealed red and gold corals 

(Parrish et al. 2002). More recent satellite tagging 

of seals at the northern extent of the Northwestern 

Hawaiian Islands shows similar feeding patterns 

to subphotic depths (Stewart et al. 2006). Monk 

seals have also been observed from the Pisces V 

submersible at more than 500 m while scientists 

were conducting coral surveys (A. Baco pers. 

obs.). The video of the encounter shows the 

seals briefly looking over the submersible and 

then using the light field from the sub to look into 

holes and cracks of the bottom. 

VI. STRESSORS ON DEEP CORAL 

COMMUNITIES

Deep coral communities within the Pacific region 

may be affected by a number of natural and 

anthropogenic stressors. Natural mortality has 

been attributed to smothering by sediments and 

by bioerosion of the substrata at the attachment 

site, which leads to toppling of colonies (Grigg 

1993). Detached colonies are rarely able to 

reattach. 

The life history attributes of deep corals makes 

them highly vulnerable to habitat damage 

associated with fishing gear and overexploitation 

in unmanaged coral fisheries (Table 4.2). Many 

year classes are exposed to effects at the same 

time. During intensive periods of indiscriminate 

fishing using bottom damaging gear, decades 

of accumulated coral growth can be lost (Grigg 

1993).

Fishing effects 

Bottom Trawling

Mobile bottom-tending gear (e.g., trawls, dredges) 

are banned in the U.S. Pacific Islands Region. 

From 1967 to 1975, Soviet and Japanese trawlers 

fished the seamounts at the south end of the 

Emperor Chain (e.g., Coco Seamount, Milwaukee 

Seamount, Colahan Seamount) and some of 

the seamounts at the north end of the Hawaiian 
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PACIFIC ISLANDS

169

Archipelago, including seamounts that were later 

designated as part of the U.S. exclusive economic 

zone (e.g., Hancock Seamount, Seamount 10, 

Seamount 11, Ladd Seamount) (Uchida and 

Tagami 1984). The primary fishing target was 

Pseudopentaceros wheeleri (armorhead) and 

Beryx spp. (alfonsinos)(Uchida and Tagami 1984). 

In 2003, submersible dives at Seamount 11 

found a large area with coral stumps and no new 

colonization (A. Baco pers. obs). The destruction 

was confined to a large swath within a fairly large 

precious coral bed (previously unknown). It could 

not be determined if this affected area was the 

result of mobile bottom-tending gear used in the 

early 1970s but if so, recovery clearly requires 

decades.

Longline fishing

Bottom longlining is not permitted in the Pacific 

Islands Region. Pelagic longlining for tuna and 

swordfish is permitted and is the region’s largest 

fishery. Longlines must be set at least 25 miles, 

and in most cases 50–75 miles from emergent 

parts of the Hawaiian Archipelago (WPFMC 

1991). This regulation was adopted to prevent 

conflicts with the coastal trolling fishery but it 

also reduces the possibility that the gear will 

affect deep corals on the slopes and seamounts 

of the Hawaiian Ridge. One exception is 

Cross Seamount located ~ 100 miles south of 

Oahu. A popular fishing site for monofilament 

handline fishing and some longline activity, it 

has accumulated numerous large fragments of 

monofilament line draped over the summit (F. 

Parrish pers. obs). Some of these line fragments 

have been seen entangled in Gerardia sp. 

colonies (A. Baco pers. obs.) and other coral 

trees appear to have been damaged. As this is 

the only location that impacts to deep coral from 

monofilament fishing have been documented the 

assigned impact rating is “Low.” 

Traps 

Bottom-set traps have been used to catch lobster 

and shrimp in the Hawaiian Archipelago. The 

Northwestern Hawaiian Islands trap fishery for 

Panulirus marginatus (Hawaiian spiny lobster) 

and Scyllarides squammosus (slipper lobster) is 

now closed, but had always operated in waters 

shallower than deep coral habitat (Polovina 

1994, Dinardo and Moffitt 2007). Trap fishing for 

the deepwater shrimp Heterocarpus laevigatus

and Heterocarpus ensifer is a small-scale pulse 

fishery limited to the main Hawaiian Islands that 

has landed 680 metric tons since the fishery’s 

inception in 1984 (PIFSC IR-06-010). The shrimp 

trapping overlaps the depth range of deep corals 

(Ralston and Tagami 1992; Moffitt and Parrish 

1992), but actual impacts to deep corals have not 

been documented. Shrimp have been observed 

associated with hard bottom features (Moffitt and 

Parrish 1992) and if fishers seek hard bottom to 

set their traps, there is potential for damage to 

deep corals. In Table 4.2, the overall gear impact 

rating of shrimp trapping was classified “Low” 

because of the small size and localized nature 

of the fishery. However if the fishery expands 

the potential impacts to deep corals would be 

an important consideration. There are no other 

recognized trap fisheries operating in the U.S. 

Western Pacific Islands and if any recreational 

or artesianal trapping is happening, it is at a very 

small scale and in shallow depths.

Other

Fishing for reef species and bottom fish typically 

rely on spearing and hook and line fishing. 

Spearfishing is largely constrained to the 

shallowest depths and is unlikely to have an effect 

on deep corals. Corals might be damaged by 

the 3-kg bottom weight used to lower handlines 

for bottom fishing or might be snagged by the 

attached hooks. However, visual surveys from 

submersibles have inspected popular bottom 

fishing sites in the main and Northwestern 

Hawaiian Islands for fishing impacts and have 

found little or no derelict gear (Kelley et al. 

2006), and there are no reports of coral bycatch 

(WPFMC 2005). 

Effects of other human activities

Coral Harvesting 

The commercial harvest of coral is the best 

documented effect to black corals and precious 

corals (pink, red, and gold) within the Hawaiian 

Archipelago. Coral harvesting has been subject 

to management under both federal and state 

regulations since the 1980s. Commercial harvest 

of black coral has always been selective, collected 

by scuba divers using hand tools. The deeper 

precious coral beds were fished initially using 

nonselective tangle net dredges, but regulations 

now require the use of selective methodologies 

such as a submersible (Figure 4.8). Commercial 

harvesting of black and precious corals has not 

been reported elsewhere in the U.S. Pacific 

outside of the Hawaiian Archipelago. 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

Maui Divers, Inc. established the small black 

coral industry in 1960 and has kept the fishery 

operating at varying levels continuously to the 

present. Limited commercial harvest of black 

coral also occurred in two other beds off Hawaii 

and Kauai. State records indicate a mean annual 

harvest of black coral at 1014 kg yr-1 from 1981 

to 1997 (Grigg 2001). Between 1999 and 2002 

there was an increase in demand accompanied 

by an increase in harvesting (Grigg 2004). This 

fishery often operates with fewer than three 

fishers so confidentiality prevents disclosure 

of annual data. Aggregating data into 7-year 

bins showed landings increased from 1985 to 

2005 with the 1999-2005 7-year bin at 22 mt 

which is more than double the prior 7-year bins 

(WPRFMC 2006). Much of this increase has been 

attributed to improved efficiency in fishing due 

to the availability of detailed bathymetric maps 

and adoption of GPS positioning. Although 11 

genera of antipatharians have been reported in 

international trade, only three species (Antipathes

cf. curvata {formerly Antipathes dichotoma} 

Antipathes grandis and Myriopathes ulex) have 

been commercially harvested in Hawaiian waters, 

with >90% of the harvested coral consisting of A. 

c.f. curvata. Other black coral species known to 

exist in this region are found in deeper waters 

and are not considered to be of commercial grade 

(Grigg 1993). 

In 1965, a bed of commercial grade pink coral 

was discovered at about 400 m depth on the 

Milwaukee Banks in the Emperor Seamount 

Chain. In 1966, Corallium secundum was also 

discovered in the Makapuu Bed off Oahu, and a 

small group of fishermen dredged the bed using 

tangle nets (Grigg 1993). Maui Divers of Hawaii 

Figure 4.8. A derelict coral dredge lost during the earliest days of the fishery (Dredge photo credit: A. Baco, 

WHOI). Inset is the Deepworker submersible which was the most recent harvesting tool employed. Photo 

credit: American Deepwater Engineering.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

171

began using a manned submersible to selectively 

harvest pink, gold, and bamboo coral from this 

bed. They removed a total of 8227 kg of pink 

coral and 2097 kg of gold coral between 1973 

and 1978 and then discontinued fishing as a 

result of high operating costs (Grigg 1993). Pink 

corals were also harvested in 1988 from Hancock 

Seamount using nonselective gear, although only 

450 kg of C. secundum were obtained, most of 

which were considered poor quality (Grigg 1993). 

In 1978, an undescribed species of Corallium

was discovered at Midway Grounds (Emperor 

Seamounts) at depths of 900–1500 m. This 

resulted in a “coral rush,” with more than 100 

boats from Japan and Taiwan operating in this 

area. Total yield exceeded 300 metric tons from 

1979 to 1984 and then dropped off because of 

resource depletion (Grigg 1993). 

In 1999, a Hawaii-based marine salvage and 

engineering company bought two deep-worker 

submersibles and began commercial harvesting 

of deep corals at the Makapuu and Keahole coral 

beds. Operations targeted pink, red, and gold 

corals. Harvesting ended in 2001, when their 

first coral auction indicated the price of the coral 

was too low to make submersible operations cost 

effective, and potential harvesting grounds in the 

Northwestern Hawaiian Islands were eliminated 

as a result of Presidential Executive Order 13196, 

which formed the Coral Reef Ecosystem Reserve 

in the Northwestern Hawaiian Islands (Grigg 

2002). Because the fishery is made up of a single 

company, confidentiality prevents reporting of 

landings data. However the permitted quota was 

not filled at either of the two beds where corals 

were harvested. Grigg (2002), working closely 

with the industry, reported removal of 60% of the 

allowed coral quota (1,216 kg) at the Makapuu 

Bed and 20% (211 kg) at the Keahole Bed. The 

precious coral fishery remains dormant today.

Illegal coral dredging

Currently, the threat of illegal coral dredging is 

thought to be remote. It is included here because 

foreign fishing vessels were documented illegally 

coral dredging in the remote Northwestern 

Hawaiian Islands in the early 1970s (Grigg 

1993). Currently, there is no evidence or even 

rumors of such illegal activity. However, much of 

the Pacific region is remote and unpopulated and 

any such activity could go undetected. Given the 

slow growth of deep corals and low recruitment 

rates, even brief periods of illegal dredging could 

have lasting effects. 

Invasive species

In 1972, the alien soft coral Carijoa riisei (Family 

Clavulariidae) was found in the fouling community 

of Pearl Harbor (Englund 2002). Originally 

thought to have colonized from the tropical 

Atlantic, recent genetic work (Samuel Kahng pers. 

comm.) suggests it arrived from elsewhere. 

It has and continues to spread to other 

suitable areas in Hawaii with high flow and 

low light (Figure 4.9). In 2001 deepwater 

surveys of the Auau Channel black coral 

beds using submarines revealed that more 

than 50% of the black coral, particularly the 

deeper, large reproductive colonies, were 

overgrown and killed by Carijoa (Kahng and 

Grigg 2005). However, Carijoa was rare on 

black coral trees in waters shallower than 

75 m (Boland and Parrish 2005). Light 

levels are thought to be too high for Carijoa

to colonize the shallower black coral trees. 

This invasive coral has been identified as a 

risk to the black coral fishery. Historically, 

black coral trees that were too deep to be 

harvested by divers were thought to serve 

as a de facto reserve for the fishery. With 

the recent discovery that many of the deep 

colonies have been killed by Carijoa, current 

management practices are being reviewed 

(Grigg 2004). Preliminary surveys of black 

Figure 4.9. The invasive gorgonian octocoral Carijoa riisei, 

that infests the deeper black corals of the Auau Channel 

beds.Photo credit: F. Parrish, NOAA Fisheries.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

coral beds on Kauai and the island of Hawaii 

have not found an infestation of Carijoa. The 

occurrence of Carijoa or other invasive species 

on deep corals elsewhere in the Pacific has not 

been reported. 

VII. MANAGEMENT OF FISHERY 

RESOURCES AND HABITATS

The Pacific Islands Region has a 25-year history 

of managing deep corals associated with its 

Precious Coral Fishery Management Plan. The 

plan was the first fishery management plan 

approved by NMFS when the Western Pacific 

Fishery Management Council was established 

as part of the national network of fishery councils 

(WPFMC 1980). The plan was implemented in 

September 1983 (48 FR 39229). 

Management of fishery resources and habitats

Historically, research has been focused on 

taxonomy (Grigg and Bayer 1976), coral 

assessments and estimates of age and growth 

to support the information needs to manage 

the fishery (Grigg 1965, 1974, 1988b, 1993, 

2001). In recent years, management research 

has expanded to include work assessing the 

connectivity of coral taxa across the Hawaiian 

Archipelago (Baco and Shank 2005; Baco et al. 

2006; Baco in prep.) and potential ecological 

associations between corals and other fauna 

(Parrish et al. 2002; Boland and Parrish 2005; 

Parrish 2006; Baco and Shirley in prep.). Studies 

are now focusing on the threat Carijoa riisei

presents to the black coral stock (Grigg 2002; 

Kahng and Grigg 2002). Replanting corals has 

been discussed as a means to mitigate impacts 

to the black coral stock from Carijoa and coral 

harvesting. Some preliminary “replanting” 

research has been conducted with Hawaiian black 

coral (Montgomery 2002), and coral harvesters 

have expressed interest in continuing the work. 

The expeditions that supported all this research 

established study sites, deployed thermographs, 

and marked colonies for future remeasurement 

to validate growth and monitor the deep coral 

ecosystem. 

Mapping Research

Future coral research will have the benefit of recent 

multibeam sonar mapping efforts. Supplementing 

earlier sidescan sonar (GLORIA system) and 

single beam sonar mapping efforts, multibeam 

products, including detailed bathymetry and 

backscatter imagery, have been made for the 

Hawaiian archipelago and other portions of the 

U.S. Pacific (Products by John Smith at HURL; 

Miller et al. 2003; Parke and Wang 2005). These 

efforts will provide a fundamental bathymetric 

context that future coral surveys will be able to use 

to infer the likelihood of deep corals. Efforts are 

currently underway to test laser-line scan survey 

technology on the black coral beds of the main 

Hawaiian Islands as a more promising means of 

directly surveying the colony abundance of deep 

corals. 

Fisheries Management Council

The Western Pacific Fishery Management Council 

(WPFMC) has responsibility for preparing fishery 

management plans (FMPs) for the fisheries in the 

U.S. exclusive economic zone (EEZ) of the Pacific 

Islands Region. Because of the steep relief of 

many Pacific Islands, deep corals also occur 

within state and territory waters, and fisheries 

can also be governed by state and territory laws 

and regulations. The Freely Associated States 

are sovereign countries and management of 

fisheries within their EEZs is governed by their 

own laws and regulations. WPFMC, an early 

leader in managing habitat impacts of fishing 

gears, prohibited demersal fish trawls, bottomset longlines, and bottom-set gillnets throughout 

the U.S. Pacific Island EEZ in 1983. The State 

and territorial laws of Hawaii, Guam, CNMI, and 

American Samoa all prohibit the use of demersal 

fish trawls within their waters. The sovereign 

territories of Kingman Reef, Palmyra Atoll, Jarvis 

Island, Howland Island, Baker Island, Midway 

Island, and Rose Atoll are National Wildlife 

Refuges administered by the U.S. Fish and 

Wildlife Service, while Wake Island and Johnston 

Atoll are managed by the Department of Defense. 

Commercial fishing is not allowed within the Fish 

and Wildlife Refuges. Thus, throughout the 

region, both shallow and deep corals have been 

largely spared impacts from trawling, at least 

within the last 25 years. 

Directed Harvest

The Precious Coral FMP and its regulations 

classify known coral beds within the western 

Pacific region and designate the harvesting 

method and amount of corals that can be 

harvested from each bed. All the known coral 

beds are in the Hawaiian Archipelago but the 

FMP includes provisions for exploratory fishing 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

173

in other areas of the U.S. Pacific. The beds are 

classified as: 1) Established Beds, 2) Conditional 

Beds, 3) Refugia Beds, and 4) Exploratory Permit 

Areas. Established beds have a history of harvest 

for which maximum sustainable yield (MSY) has 

been determined. Makapuu is the only designated 

Established Bed although the Auau black coral 

bed is in the process of being designated as 

established. Conditional beds have MSYs 

estimated based on their perceived size relative 

to established beds. There are four conditional 

beds: Keahole Point, Kaena Point, Brooks Banks 

and 180 Fathom Bank. The WESTPAC Bed is 

designated as a refugia bed, where no harvest 

is permitted. Exploratory permit areas include 

four unexplored portions of the EEZ around 

Hawaii, Guam and CNMI, American Samoa, 

and all remaining U.S. Island Possessions. The 

FMP, as amended in 2002, prohibits the use of 

nonselective gear (e.g., tangle nets, dredges) 

throughout the management area. Black coral 

is primarily found in State waters and the State 

and the WPFMC jointly manage the resource. 

Quotas and minimum size limits are monitored 

through mandatory reporting to NMFS and the 

Hawaii State Division of Aquatic Resources using 

coral landing logs and buyer reports. 

Currently, two precious coral issues are 

progressing through the WPFMC process. The 

first is reconciling coral lifespan estimates derived 

from radiometry studies (Roark et al. 2006) with 

prior estimates made from the size structure 

distribution of coral colonies and ring counts 

from basal stem cross sections. Of the three 

commercial corals, the black coral (Antipathes cf. 

curvata) radiometric estimates were consistent 

with growth rates estimated from size structure 

data (Grigg and Bayer 1976). The radiometric 

life span of pink coral was twice prior estimates, 

and gold coral (Gerardia sp.) was estimated at 

more than an order magnitude longer lived than 

prior growth estimates (Grigg 2002). This has 

prompted the WPFMC to put a 5-year moratorium 

on the fishing of gold coral until the conflicting 

lifespan data can be resolved. The second 

issue is concern that the Maui black coral bed 

may be experiencing reduced recruitment (Tony 

Montgomery, State of Hawaii, unpublished data; 

WPFMC 2006). This uncertainty combined with 

the loss of a portion of the stock to Carijoa riisei

leaves today’s biomass at least 25% lower than 

assessments in 1976 (Grigg 2004). 

Closed areas 

As noted above, all U.S. State and Federal 

waters in the Pacific Islands are closed to 

trawling and dredging—the fishing techniques 

most destructive to deep corals. Additional 

restrictions on fishing and other potentially 

harmful activities are in place in the National 

Wildlife Refuges, Papahānaumokuākea Marine 

National Monument, and in marine protected 

areas within state or territory waters.

The only area that was specifically closed to 

protect deep corals was WESTPAC Bank (located 

N.W. of Nihoa) in the Northwestern Hawaiian 

Islands. It was set aside by the precious coral 

FMP as a refuge from coral harvesting. Despite 

some interest, domestic precious coral fishing 

has never occurred in the Northwestern Hawaiian 

Islands. On determining that monk seals 

were visiting precious coral beds, the WPFMC 

proposed expanding the refuge to include areas 

where seals were visiting. Superseding this move, 

the Northwestern Hawaiian Islands Coral Reef 

Ecosystem Reserve was established in 2001 by 

Executive Order (No. 13178 and No. 13196) and 

prohibited most commercial fishing, including 

all harvesting of deep corals in the Reserve. In 

2006, while the Northwestern Hawaiian Islands 

Reserve was undergoing the designation process 

to become a national marine sanctuary, it was 

proclaimed a national monument by Presidential 

Order under the Antiquity Act of 1906 and 

renamed Papahānaumokuākea Marine National 

Monument. Within the main Hawaiian Islands 

and elsewhere in the Pacific, marine protected 

areas have not been established specifically for 

the purpose of protecting deep coral communities. 

However, there is interest from managers and 

coral harvesters to establish a closed area off 

Maui specifically for black coral to serve as a 

reproductive reserve and a biological reference 

site. 

Minerals Management Service 

Oil or gas exploration does not occur in the 

Pacific Islands Region. Historically, some 

research has focused on the prospect of 

mining manganese nodules that are formed at 

abyssal depths. Recently, interest in cobalt-rich 

manganese mining has resurged and large areas 

of the Pacific seabed, some of which include 

U.S. Pacific Islands and seamounts, are part of 

the potential mining areas (International Seabed 

Authority www.isa.org.jm/en/seabedarea/default.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

asp). Further studies of these cobalt-rich regions 

to determine deep-coral abundance and potential 

mining impacts should be considered a high 

priority.

VIII. REGIONAL PRIORITIES TO 

UNDERSTAND AND CONSERVE 

DEEP CORAL COMMUNITIES

The priorities for future work can be divided into 

the following: 1) mapping and species inventory 

of deep corals, and 2) determining the important 

physiological and ecological components of 

deep coral ecosystems. The inherently fragile 

and patchy nature of deep corals means that 

determining where they are found is a primary 

goal. Good success has been achieved using 

available mapping, remotely operated vehicles 

(ROV), and submersible infrastructure. This 

should continue and extend out to the more 

remote areas of the U.S. Pacific Islands Region. 

Also, research is needed to validate promising 

new tools to assess coral stocks with an initial 

focus of using such gear at sites previously 

surveyed using visual methods. 

1) Highest mapping and assessment priorities

 Assessments in the remote Pacific—

Historically, most research has occurred 

in and around the Hawaiian Archipelago, 

leaving American Samoa, Guam, CNMI, and 

the rest of the western Pacific unstudied. 

Baseline assessments are needed for these 

other areas, particularly those that may be 

affected by cobalt-manganese mining or 

fishery activities. Documenting areas with 

extensive coral resources will permit more 

focused enforcement and conservation 

effort. These assessments will also provide 

the DNA material for connectivity work, 

provide samples to improve taxonomy and 

systematics of deep corals, and provide an 

invaluable test to current theories on deep 

coral biogeography.

 Deeper surveys—Although the Hawaiian 

Archipelago has had some studies, few 

baseline assessments of deep corals have 

occurred outside of precious corals depth 

and none below 1800 m. There is a need to 

survey deeper habitats to better determine the 

species ranges, biodiversity, and abundance 

of deep corals. 

 Taxonomy—Critical to all of the assessment 

and ecology studies will be a dedicated effort 

to improve the taxonomy and systematics of 

deep corals, and to increase the number of 

people trained to identify these corals. There 

are very few deep coral experts in the world 

and there are currently more groups of deep 

corals needing revision and new species 

needing description than these taxonomists 

can complete in their lifetimes.

 Coral recovery studies—Beds that have 

been commercially harvested or impacted by 

fishing gear and coral harvesting should be 

periodically reassessed to determine whether 

or not the coral taxa are recovering. The 

seamounts that were subjected to bottom 

trawling or illegal harvesting more than 30 

years ago should be surveyed for signs of 

coral recovery. 

2) Physiological and ecological components of 

the deep coral ecosystem

 Environmental parameters for deep corals—

The patchy nature of deep corals, even in 

areas with similar substrate, relief, and depth 

implies that their distribution is influenced 

by other biological or environmental factors. 

Understanding the oceanographic factors 

that influence coral distributions will be 

fundamental to evaluating deep corals as a 

climate record as well as predicting where 

they might occur in unexplored areas.

 Life history, population connectivity, and 

biogeography—There is a need to understand 

more about the life history, reproduction, 

recruitment, growth, and dispersal abilities 

of deep corals; how the populations are 

connected within island/seamount chains 

and between them, and how the islands and 

seamounts of the U.S. Pacific are connected 

genetically and biogeographically to other 

parts of the Pacific. 

 Species associations—The ecological 

contribution of corals to their associated 

community needs greater attention, 

particularly the invertebrates, which are more 

likely to be dependent on the coral colonies. 

Subphotic fish communities are likely to be 

different in the remote regions of the Pacific 

and may be more tightly associated with deep 

corals.

 The ecological impact of Carijoa riisei—

Carijoa riisei represents the most clear and 

present threat to the stock of black coral and 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

175

has important implications to the black coral 

fishery. Research is needed to identify and 

understand possible ecological changes to 

the community and develop means to limit 

the spread of Carijoa riisei to other black 

coral habitats. Some research should be 

conducted to determine the feasibility of 

remediation efforts for black coral through 

replanting programs. 

IX. CONCLUSIONS

	 Deep corals occur throughout the U.S. 

Pacific but only the Hawaiian Archipelago 

and Line Islands have been the subject 

of any surveys. Coral habitat is patchy, 

suggesting at least a basic need for 

suitable bottom type and conditions of rapid 

flow. The gradients in dissolved oxygen, 

temperature, suspended particulates, etc., 

are less understood and are a priority for 

future work. Available surveys indicate 

coral beds dense with colonies that cover 

large areas are the exception. Given the 

region has little history of trawling and 

mobile bottom-tending gear, it is reasonable 

to assume this is the natural condition. 

	 Hawaiian fish are known to opportunistically 

use the corals as shelter and to some 

degree they co-occur with corals in high 

flow habitats. It is not known how the 

fish behave with deep corals in the other 

parts of the Pacific. Invertebrates are 

largely unstudied and the degree of their 

association with deep corals is unknown 

but likely to be greater. 

	 Currently, the greatest threat to corals is the 

potential for spread of the invasive species 

Carijoa riisei from the Auau channel to other 

black coral beds on Kauai and Hawaii. 

Following that, the harvesting in the 3 beds 

where the coral fishery operates needs to 

be closely monitored. For the black coral 

bed in the Auau channel, attention is needed 

because of the unanticipated loss of black 

coral to Carijoa riisei. At Makapuu, the 

regrowth of pink coral has been documented 

once and should be checked for continued 

resilience. The Keahole bed was targeted 

for red and gold coral, and its prospects 

for resilience are as yet unproven. Finally, 

the impacts to deep corals from derelict 

handlines/longlines at Cross Seamount 

and shrimp trapping in the main Hawaiian 

Islands should be assessed. 

	 Many of the new coral beds that have 

been identified in the Northwestern 

Hawaiian Islands are protected as part 

of the Papahānaumokuākea Marine 

National Monument. Being remote from 

the anthropogenic influences of the main 

Hawaiian Islands make them important 

biological reference sites for future 

research. 
PACIFIC ISLANDS

176

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

X. REFERENCES

Alcock A (1898) An account of the deep-sea 

Madreporaria collected by the Royal 

Indian marine survey ship Investigator. 

Calcutta: Trustees Indian Museum

Alcock A (1902) Report on the deep-sea 

Madreporaria of the Siboga Expedition. 

Siboga Expedition16a

Baco AR, Shank TM (2005) Population genetic 

structure of the Hawaiian precious 

coral Corallium laauense (Octocorallia: 

Corallidae) using microsatellites. Pages 

663-678 in Freiwald A, Roberts JM (eds.), 

Cold-water corals and ecosystems. 

Springer-Verlag Berlin Heidelberg

Baco AR, Clark AM, Shank TM (2006) Six 

microsatellite loci from the deep sea 

coral Corallium laauense (Octocorallia: 

Coralliidae) from the islands and 

seamounts of the Hawaiian archipelago. 

Molecular Ecology Notes 6:147-149

Baco AR, Shirley TS (In prep) Distribution and 

habitat preference of deep-sea coral 

associated invertebrates of Hawaiian 

precious corals.

Baco AR (In prep) Gene flow in the deepsea precious coral Corallium laauense

(Octocorallia: Coralliidae) from the 

islands and seamounts of the Hawaiian 

Archipelago.

Bayer FM (1951) Two new primnoid corals of 

the subfamily Calyptrophorinae. Journal 

of the Washington Academy of Sciences 

41:40-43

Bayer FM (1952) Description and re-description 

of the Hawaiian octocorals collected 

by the US Fish Commission Steamer 

“Albatross” (1.Alcyonacea, Stolonifera, 

Telstacea) Pacific Science 6:126-136

Bayer FM (1955) Contributions to the 

nomenclature, systematics, and 

morphology of the Octocorallia. 

Proceedings of the United States National 

Museum 105:207-220

Bayer FM (1956) Descriptions and re-descriptions 

of the Hawaiian Octocorals collected 

by the U.S. Fish Commission steamer 

Albatross 2. Gorgonacea: Scleraxonia. 

Pacific Science 10: 67-95

Bayer FM (1982) Some new and old species of 

the primnoid genus Callorgorgia Gray, 

with revalidation of the related genus 

Fanelia Gray (Coelenterata: Anthozoa). 

Proceedings of the Biological Society of 

Washington 95:110-160

Bayer FM (1990) A new Isidid octocoral 

(Anthozoa: Gorgonacea) from New 

Caledonia, with descriptions of other new 

species from elsewhere in the Pacific 

Ocean. Proceedings of the Biological 

Society of Washington 103:205-228

Bayer FM (1995) A new species of the 

gorgonacean genus Narella (Anthozoa: 

Octocorallia) from Hawaiian waters. 

Proceedings of the Biological Society of 

Washington 108:147-152

Bayer FM (1996) Three new species of 

precious coral (Anthozoa: Gorgonacea, 

Genus Corallium) from Pacific waters. 

Proceedings of the Biological Society of 

Washington 109:205-228

Bayer FM (1997) Narella nuttingi, a new 

gorgonacean octocoral of the family 

Primnoidae (Anthozoa) from the eastern 

Pacific. Proceedings of the Biological 

Society of Washington 110:511-519

Bayer FM, Stefani J (1988) A new species of 

Chrysogorgia (Octocorallia: Gorgonacea) 

from New Caledonia, with descriptions 

of some other species from the Western 

Pacific. Proceedings of the Biological 

Society of Washington 101:257-279

Bayer FM, Stefani J (1989) Primnoidae 

(Gorgonacea) de Nouvelle-Caledonie. 

Bulletin du Musee d’Histoire Naturelle de 

Paris 10:449-518

Berntson EA, Bayer FM, McArthur AG, France SC 

(2001) Phylogenetic relationships within 

the Octocorallia (Cnidaria: Anthozoa) 

based on nuclear 18s rRNA sequences. 

Marine Biology 138: 235-246
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

177

Boland RC, Parrish FA (2005) Description of fish 

assemblages in the black coral beds off 

Lahaina, Maui, Hawaii. Pacific Science 

59: 411-420

Brook G (1889) Report on the Antipatharia. 

Report of the Scientific Results of the 

Voyage of HMS Challenger 32:1-222. pls. 

1-15

Cairns SD (1978) Distichopora (Haplomerismos) 

anceps, a new Stylasterine coral 

(Coelenterata: Stylasterina) from 

deep water off the Hawaiian Islands. 

Micronesica 14:83-87

Cairns SD (1982) Antarctic and subantarctic 

Scleractina. Antarctic Research Series 3: 

74 pp.

Cairns SD (1984) New records of ahermatypic 

corals (Scleractinia) from the Hawaiian 

and Line Islands. Occasional Papers 

of the Bernice Pauahi Bishop Museum 

25(10):1-30

Cairns SD (1999) Cnidaria Anthozoa: DeepWater azooxanthellate Scleractinia from 

Vanuatu, and Wallis and Futuna Islands. 

Memoires du Museum National d’histoire 

Naturelle, 180:31-167

Cairns SD (2005) A revision of the Hawaiian 

Stylasteridae (Cnidaria: Hydrozoa: 

Athecata). Pacific Science 59:439-451

Cairns SD (2006) New records of azooxanthellate 

Scleractinia from the Hawaiian Islands. 

Occasional Papers of the Bernice Pauahi 

Bishop Museum 87:45-53 

Coutis PF, Middleton JH (2002) The physical and 

biological impact of a small island wake 

in the deep ocean. Deep Sea Research I 

49:1341-1361

Chave EH, Mundy BC (1994) Deep-sea benthic 

fish of the Hawaiian Archipelago, Cross 

Seamount, and Johnston Atoll. Pacific 

Science 48:367-409

Chave E H, Malahoff A (1998) In deeper waters. 

University of Hawaii Press, Honolulu, HI. 

p 125

Dana JD (1846) Zoophytes. U.S. exploring 

expedition during the years 1838-1842, 

under the command of Charles Wilkes, 

U.S.N. 7: i-iv + 1-740. Lea and Blanchard, 

Philadelphia.

Devaney DM, Eldridge LG (1977) Reef and shore 

fauna of Hawaii, Section 1: Protozoa 

through Ctenophora. Bishop Musuem 

Special Publication 64(1) p 278

DiNardo GT, Moffitt RB (2007) The northwestern 

Hawaiian Islands Lobster Fishery: A 

Targeted Slipper Lobster Fishery. In 

Lavalli KL, Spanier E (eds) The biology 

and fisheries of the slipper lobster CRC 

Press pp 243-261 

Druffel ER, Griffin MS, Witter A, Nelson E, 

Southon J, Kasgarian M, Vogel J (1995) 

Gerardia: Bristlecone pine of the deepsea? Geochima et Cosmochimica Acta 

59:5031-5036 

Duchassaing P, Michelotti J (1860) Memoire 

sur les coralliaires des Antilles. Memorie 

della Reale Accademia delle Scienze di 

Torino19: 279-365, pls. 1-10

Duncan PM (1872) On the structure and affinities 

of Guynia annulata Duncan, with remarks 

on the persistence of Palaeozoic types of 

Madreporaria. Philosophical Transactions 

of the Royal Society of London 162:29-

40

Duncan PM (1873) A description of the 

Madreporaria dredged up during 

expeditions of H.M.S. Porcupine in 

1869 and 1870. Part 1. Philosophical 

Transactions of the Royal Society of 

London 8:303-344, pls. 39-49

Duncan PM (1876) Notice of some deep-sea and 

littoral corals from the Atlantic Ocean, 

Caribbean, Indian, New Zealand, Persian 

Gulf, and Japan &c. Seas. Proceedings 

of the Zoological Society of London 

1876:428-442, pls. 38-41
PACIFIC ISLANDS

178

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

Ellis J, Solander D (1786) The natural history of the 

many curious and uncommon zoophytes, 

collected from various parts of the globe 

by the late John Ellis, systematically 

arranged by the late Daniel Solander. i-xii 

+ 1-208, pls. 1-63. Benjamin White and 

Sons; P. Elmsly, London

Englund RA (2002) The loss of native biodiversity 

and continuing nonindigenous species 

introductions in freshwater, estuarine, 

and wetland communities of Pearl Harbor, 

Oahu, Hawaiian Islands. Estuaries 

25:418-430

Esper EJC (1792) Die Pflanzenthiere. Vol. 2, pp. 

133-164, Nurnberg 

Esper EJC (1794) Fortsetzungen der 

Pflanzenthiere. Volume 1, parts 1-2, pp. 

1-64, Nurnberg

Feinstein N, Cairns SD (1998) Learning from 

the collector: a survey of azooxanthellate 

corals affixed by Xenophora (Gastropoda; 

Xenophoridae), with an analysis and 

discussion of attachment patterns. The 

Nautilus, 112:73-832

Genin A, Dayton PK, Lonsdale PF, Spiess FN 

(1986) Corals on seamount peaks provide 

evidence of current acceleration over 

deep-sea topography. Nature, 322:59-61

Goodman-Lowe G (1998) Diet of Hawaiian monk 

seal (Monachus schauinslandi) from the 

Northwestern Hawaiian Islands during 

1991-1994. Marine Biology 132:535-546

Gray JE (1866) Description of two new forms 

of gorgoniid corals. Proceedings of the 

Zoological Society of London, 1866:24-

27

Grigg RW (1965) Ecological studies of black coral 

in Hawaii. Pacific Science 19: 244-260 

Grigg RW (1974) Distribution and abundance of 

precious corals in Hawaii. Proceed. 2nd

International Coral Reef Symposium pp 

235-240

Grigg RW, Eldridge LG (1975). The commercial 

potential of precious corals in Micronesia. 

Part 1: The Mariana Islands. Sea Grant 

Publ. Guam University Marine Laboratory 

75-01 

Grigg RW, Bayer FM (1976) Present knowledge 

of the systematics and zoogeography of 

the order Gorgonacea in Hawaii. Pacific 

Science 30:167-175

Grigg RW, Opresko D (1977) Order Antipatharia 

(black corals). In: Devaney DM, Eldredge 

LG (eds) Reef and shore fauna of Hawaii. 

Vol. 1:242-261 

Grigg RW (1988a) Paleoceanography of the coral 

reefs in the Hawaiian-Emporer chain. 

Science 240:1737-1743

Grigg RW (1988b) Recruitment limitation of a deep 

benthic hard-bottom octocoral population 

in the Hawaiian Islands. Marine Ecology 

Progress Series 45:121-126

Grigg RW (1993) Precious coral fisheries of 

Hawaii and the U. S. Pacific Islands. 

Marine Fisheries Review 55:50-60

Grigg RW (2001) Black coral: history of a 

sustainable fishery in Hawaii. Pacific 

Science 55: 291-299

Grigg RW (2002) Precious corals in Hawaii: 

Discovery of a new bed and revised 

management measures for existing beds. 

Marine Fisheries Review 64:13-20

Grigg RW, Grossman EE, Earle SA, Gittings SR, 

Lott D, McDonaugh J, (2002) Drowned 

reefs and antecedent karst topography, 

Au,au Channel, S.E. Hawaiian Islands. 

Coral Reefs 21:73-82

Grigg RW (2004) Harvesting impacts and invasion 

by an alien species decrease estimates of 

black coral yield off Maui, Hawaii. Pacific 

Science 58:1-6

Hiles IL (1899) The Gorgonacea collected by 

Dr. Willey: A. Willey’s Zoological Results, 

Part 2. Cambridge.
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

179

Hoover JP (1998) Hawaii’s Sea Creatures, A 

guide to Hawaii’s marine invertebrates 

Mutual Publishing, Honolulu Hawaii p 

366 

Hubrecht AA (1885) On a new pennatulid from 

the Japanese Sea. Proceedings of the 

Zoological Society of London

Kahng SE, Grigg RW (2005) Impact of an alien 

octocoral, Carijoa riisei, on black corals in 

Hawaii. Coral Reefs 24:556-562 

Kamenkovich VM, Koshlyakov MN, Monin AS 

(1986), Synoptic eddies in the ocean. D. 

Reidel Publishing Company

Keller NB (1974) New data about some species 

of Madreporarian corals of the genus 

Flabellum. Trudy Instituta Okeanologii 

98:199-212

Kelley C R, Ikehara W (2006). The impacts of 

bottomfishing on Raita and West St. 

Rogatien Banks in the Northwestern 

Hawaiian Islands. Atoll Research Bulletin 

543:307

Kelley CR, Moffitt RB, Smith JR (2006) Mega to 

micro-scale classification and description 

of bottomfish essential fish habitat on 

four banks in the Northwestern Hawaiian 

Islands. Atoll Research Bulletin 543: 321

Kennett JP (1982) Marine Geology, PrenticeHall, Inc. p 813

Kinoshita K (1907) Vorlaufige Mitteilung uber 

einige neue japanische Primnoidkorallen. 

Annotationes zoologicae Japonenses 

6:229-237

Kinoshita K (1908) Primnoidae von Japan. Journal 

College of Science, Imperial University of 

Tokyo23: 1-74, pls. 1-6

Kinoshita K (1909) On the Primnoidae, a family 

of the Gorgonacea. Dobutsugaku zasshi 

21 (243):1-100

Kinoshita K (1913) Studien uber einige 

Chrysogorgiiden Japans. Journal 

College of Science, Imperial University of 

Tokyo33:1-47, pls. 1-3

Knauss JA (1996) Introduction to physical 

oceanography Prentice Hall p 338

Kolliker A (1870) Beitrage zur kenntnis der 

polypen. Verhandlungen der Physikalisch 

– Medicinischen Gesellschaft in Wurzburg, 

2:11-29, pls. 3-4

Kolliker A (1880) Report on the Pennatulida 

dredged by H.M.S. Challenger during the 

years 1873-1876. London

Kukenthal W (1907) Versuch einer Revision 

der Alcyonacean. 2. Die Familie der 

Nephthyiden. 3. Teil. Die Gattungen 

Eunepthya Verrill und Gersemia

Marenzelleria Zoologische Jahrbucher 

Systematik 24:317-390

Kukenthal W (1910) Zur kenntnis der gattung 

Anthomastus verr. In Doflein, F. (Ed.) 

Beitrage zur naturgeschichte ostasiens. 

math.-phys. Klasse K. bayer. Akad. 

Wissenschaft., Suppl. Bd. 1:1-16

Kükenthal W (1919) Gorgonaria. In 

Wissenschaftliche Ergebnisse der 

deutschen Tiefsee Expedition auf dem 

Dampfer “Valdivia” 1898-1899 13:1-946, 

pls. 30-89

Lesson RP (1831) Illustrations de zoologie. 

Paris.

Linnaeus C (1758) Systema naturae per regna 

tria naturae. Editio decima, reformata: 1-

824. Holmiae

Longenecker K, Dollar RA, Cahoon MK (2006) 

Increasing taxonomic resolution in dietary 

analysis of the Hawaiian monk seal. Atoll 

Research Bulletin 543:101

Miller JE, Hoeke RK, Appelgate TB, Johnson PJ, 

Smith, J, Bevacqua S (2003). Bathymetric 

Atlas of the Northwestern Hawaiian 

Islands, Draft.May 2003, National 

Oceanic and Atmospheric Administration 

and Hawaii Mapping Research Group, p 

65 
PACIFIC ISLANDS

180

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

Milne Edwards H, Haime J (1848a) Recherches 

sur les Polypiers. Mem. 1. Observations 

sur la structure et le development des 

Polypiers en general. Annales des 

Sciences Naturelles series 3, 9:37-89, pls 

4-6

Milne Edwards H, Haime J (1848b) Recherches 

sur les Polypiers. Mem. 3. Monographie 

des Eupsammides. Annales des Sciences 

Naturelles series 3, 10:65-114, pl. 1

Moffitt RB, Parrish FA, Polovina JJ (1989) 

Community structure, biomass and 

productivity of deepwater artificial reefs 

in Hawaii. Bulletin of Marine Science 

44:616-630

Moffitt RB, Parrish FA (1992) An assessment of 

the exploitable biomass of Heterocarpus 

laevigatus in the main Hawaiian Islands. 

Part 2: observations from a submersible. 

Fishery Bulletin 90:476-482

Montgomery AD (2002) The feasibility of 

transplanting black coral (Order 

Antipatharia). Hydrobiologia 471:157-164

Moseley HN (1876) Preliminary report to 

Professor Wyville Thompson on the true 

corals dredged by H.M.S. Challenger in 

deep water between the dates Dec. 30th, 

1870, and Aug. 31st, 1875. Proceedings 

of the Royal Society of London. Series B: 

Biological Sciences 24:544-569

Moseley HN (1880) Description of a new species 

of simple coral. Proceedings of the 

Zoological Society of London 1880:41-42 

Moseley HN (1881) Report on certain Hydriod, 

Alcyonaria, and Madreporarian corals 

procured during the voyage of the H.M.S. 

Challenger in the years 1873-1876. Part 

3. On the deep-sea Madrepararia. Report 

of the Scientific Results of the Voyage of 

HMS Challenger 2:127-208

Mullineaux LS, Mills SW (1997) A test of the 

larval retention hypothesis in seamountgenerated flows, Deep Sea Research I 

44:745-770

Muzik K (1978) A bioluminescent gorgonian, 

Lepidisis olapa, new species 

(Coelenterata: Octocorallia), from Hawaii. 

Bulletin of Marine Science 28:735-741

Muzik KM (1979) A systematic revision of 

the Hawaiian Paramuriceidae and 

Plexauridae (Coelenterata: Octocorallia). 

PhD Dissertation. University of Miami. p 

227

Nutting CC (1908) Descriptions of the Alcyonaria 

collected by the U.S. Bureau of Fisheries 

Steamer Albatross in the vicinity of the 

Hawaiian Islands in 1902. Proceedings 

of the United States National Museum 

34:543-601

Nutting CC (1910) The Gorgonacea of the Siboga 

expedition. VI. The Gorgonellidae. Siboga 

Expedition Monographs 13b:1-39, pls. 1-

11 

Nutting CC (1912) Descriptions of the Alcyonaria 

collected by the U.S. Fisheries Steamer 

Albatross, mainly in Japanese waters, 

during 1906. Proceedings of the United 

States National Museum 43:1-104, pls. 

1-21

Opresko DM (1974) A study of the classification 

of the Anthipatharia (Coelenterata: 

Anthozoa), with redescriptions of eleven 

species. Ph.D. Dissertation, Univ. Miami: 

1-149. Coral Gables, FL

Opresko DM (2002) Revision of the 

Antipatharia (Cnidaria: Anthozoa). 

Part II: Schizopathidae. Zoologische 

Mededelingen Leiden 76:411-442

Opresko DM (2003) Revision of the 

Antipatharia (Cnidaria:Anthozoa). 

Part III: Cladopathidae. Zoologische 

Mededelingen Leiden 77:495-536

Opresko DM (2005) New genera and species of 

Antipatharian corals (Cnidaria: Anthozoa) 

from the North Pacific. Zoologische 

Mededelingen 79-2 p129
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

181

Pallas PS (1766) Elenchus zoophytorum sistens 

generum adumbrationes generaliores 

et specierum cognitarum succinctas 

descriptions cum selectis auctorum 

synonymis. i-xvi, I 28, I 451. HagaeComitum

Parke MF, Wang HY (2005). Bathymetric 

Atlas of the main Hawaiian Islands – 

Information for Ecosystem Assessment 

and Monitoring. National Oceanic and 

Atmospheric Administration, PIFSC Data 

Report DR-05-002

Parrish FA, Boland RC (2004) Habitat and reeffish assemblages of bank summits in the 

Northwestern Hawaiian Islands. Marine 

Biology 144:1065-1073

Parrish FA, Abernathy K, Marshall GJ, Buhleier 

BM (2002). Hawaiian monk seals 

(Monachus schauinslandi) foraging in 

deep-water coral beds. Marine Mammal 

Science 18:244-258 

Parrish FA (2006). Precious corals, and subphotic 

fish assemblages. Atoll Research Bulletin 

543:425-438 

Parrish FA, Abernathy K (2006). Movements of 

monk seals relative to ecological depth 

zones in the lower northwestern Hawaiian 

Islands. Atoll Research Bulletin 543:113

Parrish FA (In press) Density and habitat of three 

deep-sea corals in the lower Hawaiian 

chain. Bulletin of Marine Science 

Polovina JJ (1994) The lobster fishery in the 

Northwestern Hawaiian Islands. In: 

Phillips BF, Cobb JS, Kittaka J (eds) Spiny 

lobster management. Blackwell Scientific 

publications Oxford pp 83-90

Pourtalès LF (1868) Contributions to the fauna 

of the Gulf Stream at great depths 

(second series). Bulletin of the Museum 

of Comparative Zoology at Harvard 

University 1:121-141

Pourtalès LF (1871) Deep-sea corals. Ill. 

Catalogue of the Museum of Comparative 

Zoology at Harvard University 4:1-93

Pourtalès LF (1878) Report on the corals of the 

“Blake” expedition. Bulletin of the Museum 

of Comparative Zoology at Harvard 

University 5:197-212 

Ralston S, Tagami DT (1992) An Assessment of 

the exploitable biomass of Heterocarpus 

laevigatus in the main Hawaiian islands. 

Part 1: Trapping surveys, depletion 

experiment, and length structure. Fishery 

Bulletin 90:494-504

Ridley SO (1884) Alcyonaria. In Report on the 

zoological collections made in the IndoPacific Ocean during the voyage of the 

H.M.S. Alert 1881-2: 327-365, pls. 36-38. 

British Museum, London. 

Roark EB, Guilderson TP, Dunbar RB, Ingram 

BL (2006) Radiocarbon based ages and 

growth rates: Hawaiian deep sea corals. 

Marine Ecology Progress Series 327:1-

14

Sars GO (1872) On some remarkable forms of 

animal life from the great deeps off the 

Norwegian coast. Christiana. Brogger 

and Christie

Schultze LS (1896) Beitrag zur systematic 

antipatharia. Abhandlungen der 

Senckenbergishcen Naturforschenden 

Gesellschaft 23: 1-40

Stewart BS, Antonelis GA, Baker JD, Yochem 

PK (2006) Foraging biogeography of 

Hawaiian monk seals in the Northwestern 

Hawaiian Islands. Atoll Research Bulletin 

543:131

Stiasny G (1947) De gorgonien-familie 

Acanthogorgiidae Kukenth. & Gorz. (Met 

bijzondere Inachtneming van het material 

der Siboga-Expeditie). Verh. kon. Nederl. 

Akad. Wetensch., Afd. Naturelle 43: 1-93, 

pls. 1-3. Amsterdam

Studer T (1894) Notes preliminaire sur les 

alcyonaires. Bulletin of the Museum 

of Comparative Zoology at Harvard 

University 25:53-69

Tchernia P (1980) Descriptive Regional 

Oceanography. Pergamon Press p 253
PACIFIC ISLANDS

182

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

Thomson JA, Henderson WD (1906) An account 

of the alcyonarians collected by the Royal 

Indian Marine Survey Ship Investigator in 

the Indian Ocean. I. The alcyonarians of 

the deep sea. XVI:10 pls. The Trustees of 

the Indian Museum, Calcutta. p 132

Thurman HV (1981) Introductory Oceanography. 

3rd ed. Charles E. Merrill Publishing 

Company. p 129

Uchida RN, Tagami DT (1984) Groundfish 

Fisheries and research in the vicinity of 

seamounts in the North Pacific Ocean. 

Marine Fisheries Review 46:1-17

van Pesch AJ (1914) The Antipatharia of the 

Siboga Expedition. Siboga Expedition 

Monographs Amsterdam, 17:1-258

Vaughan TW (1907) Recent Madreporaria of the 

Hawaiian Islands and Laysan. Bulletin. 

United States National Museum 59:1-

222, pls. 1-96 

Verrill AE (1883) Report on the Anthozoa, and 

on some additional species dredged by 

the Blake in 1877-1879, and by the U.S. 

Fish Commission steamer “Fish Hawk” 

in 1880-1882. Bulletin of the Museum 

of Comparative Zoology at Harvard 

University 11:1-72

Verrill AE (1928) Hawaiian shallow water 

Anthozoa. Bernice Pauahi Bishop 

Museum Bulletin 49:1-30 

Versluys J (1906) Die Gorgoniden der SibogaExpedition II. Die Primnoidae. dition. 

Siboga Expedition Monographs 13a:1-

187, pls. 1-10

Waller RG , Baco AR (In Press) Reproductive 

morphology of three species of deepwater precious corals from the Hawaiian 

Archipelago: Gerardia sp., Corallium 

secundum and Corallium laauense. 

Bulletin of Marine Science 

Wells JW (1954) Recent corals of the Marshall 

Islands. U.S. Geological Survey 

Professional Paper 260-I: 385-486

Wells JW (1982) Notes on Indo-Pacific 

scleractinian corals. Part 9. New corals 

from the Galapagos Islands. Pacific 

Science 36:211-220

Western Pacific Regional Fishery Management 

Council (1980). Fishery management 

plan and proposed regulations for the 

precious corals fishery of the western 

Pacific region. and Amendment 1 and 

environmental assessment for the fishery 

management plan for the precious corals 

fisheries of the western Pacific region. 

Honolulu, HI

Western Pacific Regional Fishery Management 

Council (1991) Final Environmental 

Impact Statement, Pelagic Fishery of 

the Western Pacific Region. WESTPAC 

NOAA Admin Award NAO7FC0680

Western Pacific Regional Fishery Management 

Council (2005) Final Environmental 

Impact Statement, Bottomfish and 

Seamount Groundfish Fishery of the 

Western Pacific Region. WESTPAC 

NOAA Admin Award NAO7FC0680

Western Pacific Regional Fishery Management 

Council (2006) 2006 Black coral science 

and management workshop report Apr 

18-19 48 p.

Wishner K, Levin L, Gowing M, Mullineaux 

L (1990) Involvement of the oxygen 

minimum in benthic zonation on a deep 

seamount. Nature 346:57-59

Wright EP (1885) The Alcyonaria. In: Report of 

the Scientific Results of the Voyage of 

HMS Challenger Narrative 1:689-693

Wright EP, Studer T (1889) Report on the 

Alcyonaria collected by H.M.S. Challenger. 

Zool. 31:1-314, pls. 1-43
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

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Higher Taxon Species Distribution

Depth Range 

(m) References

Phylum CnidariaClass AnthozoaSubclass HExacorallia (Zoantharia)Order ScleractiniaFamily Anthemiphylliidae Anthemiphyllia macrolobata HI Islands 369*** Cairns, 1999

Anthemiphyllia pacifica 

HI Islands & Bikini Atoll, RMI 205-296** Vaughan, 1907, Cairns, 1984

Family Caryophylliidae Anomocora sp. cf. A. fecunda HI Islands 201-271** Pourtalès, 1871, Cairns, 1984

Bourneotrochus stellulatus HI Islands 274-336** Cairns, 1984Caryophyllia atlantica HI Islands 1602** Duncan, 1873, Cairns, 1984Caryophyllia hawaiiensis HI Islands 44-388** Vaughan, 1907, Cairns, 1984Caryophyllia marmorea HI Islands 331-337** Cairns, 1984Caryophyllia octopali HI Islands 457-627** Vaughan, 1907, Cairns, 1984Caryophyllia rugosa HI Islands 137-439** Moseley, 1881, Cairns, 1984Caryophyllia sp. cf. C. ambrosia HI Islands 56-206** Alcock, 1898, Cairns, 1984”Ceratotrochus” laxus HI Islands 583-678** Vaughan, 1907Coenosmilia inordinata HI Islands 244-322** Cairns, 1984Conotrochus funicolumna HI Islands 165-600** Alcock, 1902, Cairns, 1984Crispatotrochus rubescens 

HI Islands & Christmas Island, Line Islands 197-634** Moseley, 1881, Cairns, 1984

Deltocyathus sp. cf. D. andamanicus HI Islands 274-518** Alcock, 1898, Cairns, 1984

Appendix 4.1. List of known species of ceep corals from the U.S. Pacific Islands. All species listed are found in Hawaii except one octocoral, Keroeides koreni and several antipatharins as noted. List for octocorals and scleractinians in Hawaii based primarily on unpublished list compiled by Dr. Stephen Cairns, Smithsonian Institution, with additions from recent Pisces cruises led by A. Baco and additions for non-Hawaiian islands based on cited literature. List for antipatharians from Hawaii and Guam unpublished list compiled by and courtesy of Dr. Dennis Opresko. * = depth range known from full range for species, including outside chapter region** = depth range known from 2 or more specimens from Hawaii, Christmas, Line Islands, etc*** = depth from a single individual from Hawaii, usually the holotype
PACIFIC ISLANDS

184

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION Higher Taxon Species Distribution Depth Range (m References Desmophyllum dianthus HI Islands Esper, 1794 Paracyathus molokensis HI Islands 161-260** Vaughan, 1907 Trochocyathus aithoseptatus HI Islands 371-454** Cairns, 1984 Trochocyathus burchae HI Islands 64*** Cairns, 1984 Trochocyathus gardineri HI Islands 274-470** Vaughan, 1907, Cairns, 1984 Trochocyathus mauiensis HI Islands 174-278** Vaughan, 1907 Trochocyathus oahensis HI Islands 75-571** Vaughan, 1907, Cairns, 1984 Trochocyathus patelliformis HI Islands 1020*** Cairns, 1999 Trochocyathus rhombocolumna HI Islands 110-530* Alcock, 1902 Family Dendrophylliidae Balanophyllia desmophyllioides HI Islands 143-406** Vaughan, 1907 Balanophyllia diomedeae HI Islands 110-307** Vaughan, 1907, Cairns, 1984 Balanophyllia gigas HI Islands 90-640* Moseley, 1881 Balanophyllia laysanensis HI Islands 238-271** Vaughan, 1907 Cladopsammia echinata HI Islands 295-470** Cairns, 1984 Cladopsammia eguchii HI Islands Wells, 1982 Eguchipsammia gaditana HI Islands 244-470** Duncan, 1873, Cairns 1984 Eguchipsammia fistula HI Islands Alcock, 1902 Eguchipsammia serpentina HI Islands 269-362** Vaughan, 1907

Enallopsammia rostrata HI Islands 362-583** Pourtalès, 1878, Cairns, 1984Endopachys grayi HI Islands 37-274**

Milne-Edwards & Haime, 1848a & b, Cairns, 1984

Family Flabellidae Flabellum marcus 

HI Islands & NW of Wake Island 1261-1602** Keller, 1974, Cairns, 1984

Flabellum pavoninum HI Islands 183-517** Lesson, 1831, Cairns, 1984Flabellum vaughani HI Islands 232-369** Cairns, 1984Javania exserta HI Islands 400*** Cairns, 2006Javania fuscus HI Islands 13-271** Vaughan, 1907Javania insignis 

HI Islands & Christmas Island, Line Islands 52-825** Duncan, 1876, Cairns, 1984
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

185

Higher Taxon Species Distribution

Depth Range 

(m References

Javania lamprotichum 

HI Islands & Johnston Atoll 244-322** Moseley, 1880, Cairns, 1984

Placotrochides minuta HI Islands 119-291*** Feinstein and Cairns, 1998, Cairns 2006Polymyces wellsi HI Islands 440-858** Cairns, 2006

Family Fungiacyathidae Fungiacyathus fissilis HI Islands 212-503** Cairns, 1984

Fungiacyathus fragilis HI Islands 1762-2056** Sars, 1872, Cairns, 1984

Family Gardineriidae Gardineria hawaiiensis HI Islands 369-541** Vaughan, 1907, Cairns, 1984, 2006Family Guyniidae Guynia annulata HI Islands 64-384** Duncan, 1872, Cairns, 1984Family Micrabaciidae Letepsammia formosissima HI Islands 109-470** Moseley, 1876, Cairns, 1984Family Pocilloporidae Madracis kauaiensis HI Islands 362-538** Vaughan 1907, Cairns 2006Family Oculinidae Madrepora kauaiensis HI Islands 362-538** Vaughan, 1907, Cairns, 1984

Madrepora oculata HI Islands 627-750** Cairns, 1984

Family Turbinoliidae Deltocyathoides orientalis HI Islands 439-494** Duncan, 1876, Cairns 1984, Cairns 2006Family Stenocyathidae Stenocyathus vermiformis 

HI Islands & S. Pacific Seamounts 439** Pourtalès, 1868, Cairns, 1982, 

Cairns, 1984

Order AntipathariaFamily Antipathidae Antipathes grandis HI Islands Verrill, 1928

Antipathes sp., cf. A. curvata HI Islands van Pesch, 1914Antipathes n. sp., cf. A. dichotoma HI Islands Pallas, 1766Antipathes sp. cf. A. flabellum Guam Pallas, 1766Antipathes sp., cf. 

A. spinuilosa Guam Schultze, 1896

Antipathes intermedia HI Islands 347-366*** Brook, 1889, Grigg and Opresko, 1977Cirrhipathes anguina HI Islands 25-40*** Dana, 1846, Grigg and Opresko, 1977Cirrhipathes contorta GuamCirrhipathes propinqua GuamCirrhipathes spiralis HI Islands Linnaeus, 1758, Grigg and Eldridge, 1975
PACIFIC ISLANDS

186

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION Higher Taxon Species Distribution Depth Range (m References Stichopathes echinulata HI Islands 305-565*** Brook, 1889, Grigg and Opresko, 1977 Family Aphanipathidae Aphanipathes sarothamnoides Guam Brook, 1889 Acanthopathes undulata HI Islands 110-490** van Pesch, 1914, Grigg and Opresko, 1977 Family Cladopathidae Trissopathes pseudotristicha HI Islands & Channel Islands 326-4539** Opresko, 2003 Trissopathes tetracrada HI Islands 375-425** Opresko, 2003 Family Leiopathidae Leiopathes glaberrima HI Islands Esper, 1792, Opresko, 1974 Leiopathes n. sp. HI Islands 403-471** Prelim. ID by Opresko, Pisces Cruise 2003 &2004 Family Myriopathidae Myriopathes ulex HI Islands & Guam Ellis and Solander, 1786, Grigg and Eldridge, 1975 Myriopathes sp., cf. M. japonica HI Islands Brook, 1889 Cupressopathes abies Guam Linnaeus, 1758 Antipathella sp., cf. A. subpinnata HI Islands 455-460*** Ellis and Solander, 1786, Grigg and Opresko, 1977 Family Schizopathidae Bathypathes alternata HI Islands 1195-1744** Brook, 1889, Pisces Cruise 2003 Bathypathes conferta HI Islands 380*** Brook, 1889, Grigg and Opresko, 1977 Bathypathes patula HI Islands Brook, 1889, Unpubl record at USNM

Stauropathes staurocrada

HI Islands & Johnston Atoll

220-441, 1400-1700** Opresko, 2002

Stauropathes sp. HI Islands 604***

Prelim. ID by Opresko, Pisces Cruise 2003

Umbellapathes helioanthes HI Islands 1205-1383** Opresko 2005Umbellapathes, new species B HI Islands 742-744***

Prelim. ID by Opresko, Pisces cruise 2004

Dendropathes bacotaylorae HI Islands 408*** Opresko 2005

Order Zoanthidea

Zoanthid blue HI Islands 352-415** Chave and Malahoff, 1998
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

187

Higher Taxon Species Distribution

Depth Range 

(m References

Zoanthid tan HI Islands 500-1910** Chave and Malahoff, 1998Gerardia sp. HI Islands 343-577**

Ryland and Baco in prep., Chave and Malahoff, 1998

Parazoanthus sp. 1 HI Islands 343-460** Chave and Malahoff, 1998Parazoanthus sp. 2 HI Islands 332-1025** Chave and Malahoff, 1998

Subclass OctocoralliaOrder AlcyonaceaFamily Alcyoniidae Anthomastus fisheri HI Islands 356-462** Bayer, 1952, Chave and Malahoff, 1998

Anthomastus (Bathyalcyon) robustus HI Islands Versluys, 1906, fide Williams Anthomastus granulosus HI Islands 20-201** Kukenthal 1910, Bayer, 1952Inflatocalyx sp. HI Islands fide Williams

Family Clavulariidae Carijoa riisei Invasive, HI Islands Duch. And Mich., 1860

Clavularia grandiflora HI Islands 966* Nutting, 1908, Bayer, 1952Telestula corrugata HI Islands 914* Nutting, 1908, Bayer, 1952Telestula spiculicola HI Islands 518-616* Nutting, 1908, Bayer, 1952Telestula spiculicola robusta HI Islands Bayer, 1952

Family Nidaliidae Nidalia sp. HI Islands fide Williams

Siphonogorgia alexanderi HI Islands 223-283* Nutting, 1908, Bayer, 1952Siphonogorgia collaris HI Islands 144* Nutting, 1908, Bayer, 1952

Order GorgonaceaFamily Acanthogorgiidae Acanthogorgia sp. cf. A. striata Nutting, 1911 HI Islands 215-564** Grigg and Bayer, 1976

Acanthogorgia n. sp. HI Islands Muzik, 1979Acanthogorgia sp. cf. A. paramuricata HI Islands 350-396** Stiasny, 1947, Grigg and Bayer, 1976 Acanthogorgia sp. HI Islands 1295*** Berntson et al., 2001 Cyclomuricea flabellata HI Islands 71-396** Nutting, 1908, Grigg and Bayer, 1976Muricella tenera HI Islands 237-2533* Ridley, 1884, Nutting, 1908

Family Anthothelidae Anthothela nuttingi HI Islands

340-465, 1387-1820** Bayer, 1956, Grigg and Bayer, 1976
PACIFIC ISLANDS

188

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION Higher Taxon Species Distribution Depth Range (m References Antothela n. sp. 1 HI Islands 1319*** Prelim. ID by Cairns, Pisces Cruise 2003 Antothela n. sp. 2 HI Islands 1804*** Prelim. ID by Cairns, Pisces Cruise 2003 Family Chrysogorgiidae Chrysogorgia arborescens HI Islands 722-914* Nutting, 1908 Chrysogorgia chryseis HI Islands 732*** Bayer and Stefani, 1988, Berntson et al., 2001 Chrysogorgia delicata HI Islands 536-1463* Nutting, 1908 Chrysogorgia elegans HI Islands 433-634* Verrill, 1883, Nutting, 1908 Chrysogorgia flavescens HI Islands 1688-1977* Nutting, 1908 Chrysogorgia geniculata HI Islands 146-616* Wright & Studer, 1889, Nutting, 1908 Chrysogorgia sp. cf. C. japonica HI Islands 750-1050** Wright & Studer, 1889, Grigg and Bayer, 1976 Chrysogorgia paillosa HI Islands 704-1858* Kinoshita, 1913, Grigg and Bayer, 1976, Nutting, 1908 Chrysogorgia scintillans HI Islands 580-2050** Bayer and Stefani, 1988, Chave and Malahoff, 1998 Chrysogorgia stellata HI Islands 649-678* Nutting, 1908 Chrysogorgia sp. cf. C. stellata HI Islands 646-675, 830- 922* Nutting, 1908, Grigg and Bayer, 1976, Bayer and Stefani 1988 Chrysogorgia n. sp. (1/3R) HI Islands 1204*** Prelim. ID by Cairns, Pisces Cruise 2003

Chrysogorgia n. sp (2/5L) HI Islands 691-742** Prelim. ID by Cairns, Pisces Cruise 2004Iridogorgia superba HI Islands 704-914* Nutting, 1908, Grigg and Bayer, 1976Iridogorgia bella HI Islands 750-1925** Nutting, 1908, Chave and Malahoff, 1998Iridogorgia. n. sp. HI Islands 1443*** Prelim. ID by Cairns, Pisces Cruise 2003Metallogorgia melanotrichos HI Islands 183-1385* Wright and Studer, 1889, Nutting, 1908Metallogorgia n. sp. HI Islands 1805*** Prelim. ID by Cairns, Pisces Cruise 2003Pleurogorgia militaris HI Islands 2142* Nutting, 1908Radicipes spiralis HI Islands 258** Nutting, 1908, Grigg and Bayer, 1976
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

189

Higher Taxon Species Distribution

Depth Range 

(m References

Family Coralliidae Corallium abyssale HI Islands 1829-2403*** Bayer, 1956, Bayer unpubl ms

Corallium ducale HI Islands Bayer, 1955, Bayer unpubl msCorallium kishinouyei HI Islands 1145*** Bayer, 1996, Bayer unpubl msCorallium laauense HI Islands 365-580** Bayer, 1956, Grigg and Bayer, 1976Corallium niveum HI Islands 232-282*** Bayer, 1956, Bayer unpubl msCorallium regale HI Islands 365-719** Bayer, 1956, Grigg and Bayer, 1976Corallium secundum HI Islands 231-576**

Dana, 1846, Bayer, 1956, Grigg and Bayer, 1976, Pisces Cruise 2003

Corallium imperiale HI Islands 1096*** Bayer, 1955, Pisces Cruise 2003  Corallium cf. secundum HI Islands Prelim. ID by Cairns, Pisces Cruise 2003Corallium laauense x halmahera HI Islands Prelim. ID by Cairns, Pisces Cruise 2003 Corallium n. sp. HI Islands Prelim. ID by Cairns, Pisces Cruise 2003Paracorallium tortuosum HI Islands 167-408** Bayer, 1956, Grigg and Bayer, 1976

Family Gorgoniidae Eunicella n. sp. 

A HI Islands 275-495** Grigg and Bayer, 1976 

Family Isididae Acanella dispar HI Islands 275-445** Bayer, 1990

Acanella weberi HI Islands Nutting, 1910Isidella trichotoma HI Islands 1920*** Bayer, 1990Isidella sp. “5” HI Islands Muzik museum idIsidella n. sp. (lyrate) HI Islands 1808*** Prelim. ID by Cairns, Pisces Cruise 2003Keratoisis flabellum HI Islands 346-465** Nutting, 1908, Grigg and Bayer, 1976Keratoisis grandis HI Islands 1344-1582* Nutting, 1908Keratoisis n. sp. HI Islands 305-565** Grigg and Bayer, 1976Lepidisis nuda HI Islands

Wright and Studer, 1889, Grigg and Bayer 1976

Lepidisis olapa HI Islands 215-665** Muzik, 1978Lepidisis paucispinosa HI Islands 539-631**

Wright and Studer, 1889, Nutting 1908, Muzik, 1978

Lepidisis sp. HI Islands 1425*** Berntson et al., 2001
PACIFIC ISLANDS

190

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION Higher Taxon Species Distribution Depth Range (m References Family Keroeididae Keroeides fallax HI Islands 238-245*** Bayer, 1956 Keroeides koreni Marshall Islands Wright & Studer, 1889, Bayer, 1956 Keroeides mosaica HI Islands 167-465** Bayer, 1956, Grigg and Bayer, 1976 Keroeides pallida HI Islands 146*** Hiles, 1899, Bayer, 1956 Family Paragorgiidae Paragorgia dendroides HI Islands 490-1910** Bayer, 1956, Chave and Malahoff, 1998 Paragorgia sp. cf. P. regalis Nutting, 1912 HI Islands 350-396** Grigg and Bayer, 1976 Paragorgia n. sp. HI Islands 350-396** Grigg and Bayer, 1976 Family Plexauridae Anthomuricea tenuispina HI Islands 428-531, 581- 688** Nutting, 1908, Grigg and Bayer, 1976 Anthomuricea sp. cf. A. divergens HI Islands 381-426** Kükenthal, 1919, Grigg and Bayer, 1976 Anthomuricea sp. cf. A. reticulata HI Islands 362-421** Nutting, 1910, Grigg and Bayer, 1976 Anthomuricea n. sp. A HI Islands Muzik, 1979 Bebryce brunnea HI Islands 167-396** Nutting, 1908, Grigg and Bayer, 1976 Bebryce n. sp. HI Islands Muzik, 1979 Muriceides sp. A HI Islands Muzik, 1979 Muriceides sp. B HI Islands Muzik, 1979 Filigella n. sp. Thesea n. sp. HI Islands Muzik, 1979 Muriceides tenuis HI Islands 232-362* Nutting, 1908, Muzik, 1979

Muriceides n. sp. A HI Islands Muzik, 1979Muriceides n. sp. B HI Islands Muzik, 1979New genus, n. sp. HI Islands Muzik, 1979Paracis horrida HI Islands

Thomson & Henderson, 1906, Muzik, 1979

Paracis miyajimai HI Islands 362-531** Kinoshita, 1909, Grigg and Bayer, 1976 Paracis n. sp. A HI Islands Muzik, 1979Paracis spinifera HI Islands 350-396** Nutting, 1912, Grigg and Bayer, 1976 Paramuricea HI Islandsensis HI Islands

350-396, 924-1241** Nutting, 1908, Grigg and Bayer, 1976 
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

PACIFIC ISLANDS

191

Higher Taxon Species Distribution

Depth Range 

(m References

Placogorgia n. sp. HI Islands Bayer unpubl msPlacogorgia sp. n. A HI Islands Muzik, 1979Placogorgia n. sp. B HI Islands Muzik, 1979Placogorgia sp. cf. P. dendritica HI Islands 350-396** Nutting, 1910, Grigg and Bayer, 1976 Placogorgia sp. HI Islands 335-375** Grigg and Bayer, 1976Placogorgia sp. cf. Ps. placoderma HI Islands 73, 182** Nutting, 1910, Grigg and Bayer, 1976 Pseudothesea sp. cf. Ps. orientalis HI Islands 147, 350-396**

Thom & Hend, 1906, Grigg and Bayer, 1976 

Swiftia n. sp. 1 HI Islands 340-365** Grigg and Bayer, 1976 Swiftia n. sp. 2 HI Islands 350-396** Grigg and Bayer, 1976 Swiftia pacifica HI Islands Muzik 1979Thesea sp. cf. T. ramosa HI Islands 313-399** Nutting, ??, Grigg and Bayer, 1976 Villogorgia arbuscula HI Islands 315-412**

Wright & Studer, 1889, Grigg and Bayer, 1976 

Villogorgia n. sp. 1 HI Islands 350-396** Grigg and Bayer, 1976 Villogorgia n. sp. 2 HI Islands 350-396** Grigg and Bayer, 1976 Villogorgia n. sp. A HI Islands Muzik, 1979Villogorgia n. sp. B HI Islands Muzik, 1979Villogorgia n. sp. C HI Islands Muzik, 1979

Family Primnoidae Callogorgia formosa HI Islands Kukenthal, 1907, Bayer, 1982

Callogorgia gilberti HI Islands 215-960** Nutting, 1908, Grigg and Bayer, 1976Callogorgia n. sp. HI Islands 350-396** Grigg and Bayer, 1976Calyptrophora agassizii HI Islands 781-1145** Studer, 1894, Grigg and Bayer, 1976Calyptrophora angularis HI Islands 1207-3292* Nutting, 1908, Grigg and Bayer, 1976Calyptrophora clarki HI Islands 12-1275 Bayer, 1951Calyptrophora japonica HI Islands 216-432** Gray, 1866, Grigg and Bayer, 1976Calyptrophora n. sp. HI Islands 344-454** Grigg and Bayer, 1976
PACIFIC ISLANDS 192

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION Higher Taxon Species Distribution Depth Range (m References Calyptrophora wyvillei HI Islands 744-823* Wright, 1885, Nutting, 1908 Calyptrophora n. sp. 1 (lyrate) HI Islands 1078*** Prelim. ID by Cairns, Pisces Cruise 2003 Calyptrophora n. sp. 2 HI Islands 1807*** Prelim. ID by Cairns, Pisces Cruise 2003 Candidella gigantea HI Islands 1720-1815** Wright & Studer, 1889, Pisces cruise 2003 Candidella helminthophora HI Islands 38-1820** Nutting, 1908, Grigg and Bayer, 1976 Fanellia euthyeia HI Islands Bayer and Stefani, 1989 Fanellia medialis HI Islands Bayer and Stefani, 1989 Fanellia tuberculata HI Islands Versluys, 1906, Bayer, 1982 Narella bowersi HI Islands 1344-1937* Nutting, 1908, Berntson et al., 2001, Grigg and Bayer, 1976 Narella dichotoma HI Islands Versluys, 1906, Bayer ms: 27 Narella sp. cf. N. megalepis HI Islands 215-564** Kinoshita, 1908, Grigg and Bayer, 1976 Narella nuttingi HI Islands 1350*** Bayer, 1997, Berntson et al., 2001 Narella ornata HI Islands 748-1007*** Bayer, 1995 Narella studeri HI Islands Versluys, 1906 Narella n. sp. 1 HI Islands 350-396** Grigg and Bayer, 1976 Narella n. sp. 2 HI Islands 353-417** Grigg and Bayer, 1976

Narella. n. sp. (unbranched) HI Islands Prelim. ID by Cairns Pisces Cruise 2003Paracalyptrophora n. sp. HI Islands 367-398** Prelim. ID by Cairns Pisces Cruise 2004Parastenella n. sp. HI Islands 517*** Prelim. ID by Cairns Pisces Cruise 2004Plumarella n. sp. HI Islands 384-432** Grigg and Bayer, 1976Thouarella (A.) biserialis HI Islands 439* Nutting, 1908, Grigg and Bayer, 1976Thouarella (A.) regularis HI Islands 183-722* Wright and Studer, 1889, Nutting, 1908Thouarella sp. cf. T. (T.) typica HI Islands 350-396** Kinoshita, 1907, Grigg and Bayer, 1976
STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION

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193

Higher Taxon Species Distribution

Depth Range 

(m References

Order PennatulaceaFamily Anthoptilidae Anthoptilum murrayi HI Islands 426-2286* Kolliker, 1880, Nutting, 1908Family Chunellidae Calibelemnon symmetricum HI Islands 196-1650** Nutting, 1908, Chave and Malahoff, 1998Family Echinoptilidae Echinoptilum macintoshi HI Islands 225-232* Hubrecht, 1885, Nutting, 1908Family Funiculinidae Funiculina sp. HI Islands 254-1940** Chave and Malahoff, 1998Family Halipteridae Halipterus willemoesi HI Islands fide WilliamsFamily Kophobelemnidae Kophobelemnon sp. (short stemmed) HI Islands Prelim. ID by Cairns, Pisces Cruise 2003Family Pennatulidae Pennatula flava HI Islands 223-316* Nutting, 1908

Pennatula pallida HI Islands 402-530* Nutting, 1908Pennatula pearceyi HI Islands 1033* Kolliker, 1880, Nutting, 1908Pennatula sanguinea HI Islands 903-1033* Nutting, 1908

Family Protoptilidae Protoptilum wrighti HI Islands 523* Nutting, 1908

Protoptilum attenuatum HI Islands 925* Nutting, 1908Protoptilum studeri HI Islands 97-421* Nutting, 1908

Family Umbellulidae Umbellula carpenteri HI Islands 1046-2056* Kolliker, 1880, Nutting 1908

Umbellula gilberti HI Islands 708-1951* Nutting, 1908Umbellula jordani HI Islands 704-2403* Nutting, 1908

Family Virgulariidae Virgularia abies HI Islands 223* Kolliker, 1870

Virgularia molle HI Islands 1265-1280* Kolliker, 1880

Class HydrozoaOrder StylasterinaFamily Stylasteridae Distichopora (Haplomerismos) anceps HI Islands 360-577** Cairns, 1978, 2005

Distichopora asulcata HI Islands 293-377** Cairns, 2005Stylaster griggi HI Islands 322-583** Cairns, 2005Stylaster infundibuliferus HI Islands 521-563** Cairns, 2005
PACIFIC ISLANDS 194

STATE OF DEEP CORAL ECOSYSTEMS IN THE PACIFIC ISLANDS REGION
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

NORTHEAST

195

STATE OF DEEP CORAL ECOSYSTEMS IN THE

NORTHEASTERN US REGION:

MAINE TO CAPE HATTERAS

David B. Packer1, Deirdre Boelke2, Vince Guida1, and Leslie-Ann McGee2

1

NOAA Fisheries Service, Northeast Fisheries 

Science Center,

James J. Howard Marine Sciences Laboratory,

Highlands, NJ 07732

2

New England Fishery Management Council,

50 Water Street, Mill 2, Newburyport, MA 

01950

I. INTRODUCTION

The U.S. Northeast Shelf Large Marine Ecosystem 

encompasses 260,000 square km and extends 

from the Gulf of Maine and Georges Bank 

southward through southern New England waters 

and the Middle Atlantic Bight to Cape Hatteras. It 

extends from the coast eastward to the edge of 

the continental shelf and slope and offshore to 

the Gulf Stream (Sherman et al. 1996). Some of 

the specific locations with known occurrences of 

deep corals include parts of the Gulf of Maine, 

Georges Bank, as well as a number of canyons 

that bisect the continental shelf and slope. Deep 

corals have also recently been collected from the 

New England Seamount chain; the seamounts 

are located off the continental shelf, rising above 

the abyssal plain, and they encompass more than 

30 major volcanic peaks extending from Georges 

Bank southeast for about 1,100 km to the eastern 

end of the Bermuda Rise (Figure 5.1). Several 

recent surveys have taken place here, although 

most of the seamount chain is located outside the 

200 nautical mile U.S. Exclusive Economic Zone 

(EEZ); however, Bear, Physalia, Mytilus, and 

Retriever Seamounts do occur within the EEZ.

In the U.S. Northeast Shelf Large Marine 

Ecosystem, deep corals have been noted since 

the surveys of Verrill in the 19th century (Verrill 

1862, 1878a, 1878b, 1879, 1884) and as fisheries 

bycatch since that period. They also occur off 

Atlantic Canada on the continental slope and in 

submarine canyons, and are particularly abundant 

in the Northeast Channel (Verrill 1922; Deichman 

1936; Breeze et al. 1997; MacIsaac et al. 2001; 

Mortensen and Buhl-Mortensen 2004; Gass and 

Martin Willison 2005; Mortensen et al. 2005). 

Many gorgonian corals such as Paragorgia have 

been regularly encountered by fishermen on 

hard substrate such as boulders, gravel, or rocky 

outcrops (e.g., Breeze et al. 1997; Leverette and 

Metaxas 2005).

This chapter summarizes the current knowledge 

of deep corals and deep coral communities for the 

U.S. Northeast Shelf Large Marine Ecosystem, 

including the diversity of coral species and their 

distribution, associated species, and habitat 

preferences. In addition, a summary of current 

stressors affecting these habitats, and existing 

conservation and management activities are 

presented. Gaps in our understanding of deep 

coral communities and a summary of research 

priorities that could help fill these gaps are 

highlighted as a key need to assist in identifying 

future actions to conserve and manage these 

vulnerable ecosystems.

However, it should be noted that to our knowledge 

the northeast region does not have the abundance 

of large, structure-forming deep corals and deep 

coral habitats that are present in other regions. 

Thus, we will confine our discussions to the 

major gorgonian species such as, for example, 

Paragorgia arborea, Primnoa resedaeformis, 

Acanthogorgia armata, and Paramuricea 

grandis, and others, as well as some of the 

more noteworthy alcyonaceans (soft corals) and 

scleractinians (stony corals).

II. GEOLOGICAL SETTING

This brief review of the pertinent geological 

characteristics of the regional systems of the 

U.S. Northeast Shelf Large Marine Ecosystem, 

as well as the subsequent section on the 

oceanographic characteristics, is based on 

several summary reviews. Literature citations 

are not included because these are generally 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

Figure 5.1 Coral Distribution - Deep coral data from the Theroux and Wigley (1998) and the Watling et al. 

(2003) databases represent known locations of stony, gorgonian, and true soft deep coral species from data 

ranging from (but not inclusive of) 1870 to the 1990s. This information does not represent all deep coral locations in this region as the U.S. Exclusive Economic Zone has not been extensively surveyed for deep corals. 

Also, certain deep coral points may no longer be present due to natural and human induced impacts that have 

occurred since the point was sampled. Essential Fish Habitat (EFH) Closed Areas prohibit the use of mobile 

bottom-tending gear indefinitely. Northeast Multispecies Closed Areas,indicated in the legend by “Closures to 

All Gear Capable of Catching Groundfish” prohibit the use of the gear capable of catching groundfish except 

in portions of the closed areas defined in the Special Access Program during certain times of the year (see 

NEFMC website). No fishing restrictions exist within the boundaries of the Stellwagen Bank NMS with the 

exception of the sliver that overlaps both the NEFMC Western Gulf of Maine Groundfish and Western Gulf 

of Maine Essential Fish Habitat Closed Areas. In addition to the EFH Closed Areas on Georges Bank and in 

southern New England, Lydonia and Oceanographer Canyons on the continental shelf-break are also considered EFH Closures and were implemented in the Monkfish FMP in 2005. Within these areas, trawling and 

gillnetting while on a monkfish (Lophius americanus, goosefish) days-at-sea (DAS) is prohibited indefinitely to 

protect the sensitive habitats therein, including deep coral and other structure-forming organisms. 
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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accepted descriptions of the regional systems. 

Source references include, and more information 

can be found in: Backus and Bourne (1987), 

Schmitz et al. (1987), Tucholke (1987), Wiebe 

et al. (1987), Cook (1988), Stumpf and Biggs 

(1988), Townsend (1992), Conkling (1995), 

Brooks (1996), Sherman et al. (1996), Beardsley 

et al. (1997), Dorsey (1998), Packer (2003), 

Stevenson et al. (2004), Auster et al. (2005), and 

Babb (2005).

Gulf of Maine

The Gulf of Maine is a semi-enclosed continental 

shelf sea of 90,700 sq. km bounded on the east by 

Browns Bank, on the north by the Scotian Shelf, 

on the west by the New England states, and on 

the south by Cape Cod and Georges Bank. It is 

distinct from the Atlantic, an ecologically separate 

sea within a sea. The Gulf is characterized by 

a system of 21 deep basins, moraines, rocky 

ledges, and banks, with limited access to the 

open ocean. Sediments in the Gulf of Maine 

are highly variable and, when coupled with the 

vertical variation of water properties found in the 

Gulf, results in a great diversity of benthic habitat 

types and benthic organisms. Sand, silt, and 

clay are found throughout the Gulf, with the finer 

sediments generally found in the deeper basins. 

Rocky substrates (which include gravel, pebbles, 

cobbles, and boulders) are found primarily in 

the Northeast Channel, with other smaller, more 

variable rocky areas interspersed in the Gulf. 

Rocky outcrops form significant features, such 

as Cashes Ledge, and benthic fauna found on 

these include sponges, tunicates, bryozoans, 

and hydroids.

Of the 21 deep basins, Jordan (190 m), Wilkinson 

(190 m), and Georges Basins (377 m), are the 

largest basins and deepest habitats within the 

Gulf of Maine. Their great depths resulted from 

glacial erosion of relatively soft rocks. The bottom 

sediments of these deep basins are generally very 

fine, featureless muds, but some gravel may also 

be found; little or no sediment transport occurs 

here. Unique invertebrate communities are found 

on the seafloor, including deep corals, ophiuroids 

(brittle stars), tube building amphipods, burrowing 

anemones, and polychaete worms.

Georges Bank

Georges Bank is a shallow (3-150 m depth), 

elongate extension of the continental shelf 

that extends from Cape Cod, Massachusetts 

(Nantucket Shoals) to Nova Scotia (Scotian 

Shelf) and covers more than 40,000 km2. It is 

characterized by a steep slope on its northern 

edge and a broad, flat, gently sloping southern 

flank. It is separated from the rest of the 

continental shelf to the west by the Great South 

Channel. The central region of the Bank is 

shallow. Bottom topography on the eastern Bank 

is characterized by linear ridges in the western 

shoal areas; relatively smooth, gently dipping 

sea floor on the deeper, eastern most part, and 

steeper and smoother topography incised by 

submarine canyons on the southeastern margin. 

The sediments vary widely, but are mostly sandy 

or silty, with coarse gravel and boulders at the 

northern margins.

At least 70 large submarine canyons occur near 

the shelf break along Georges Bank and the MidAtlantic down to Cape Hatteras, cutting into the 

slope and occasionally up into the shelf as well. 

The canyons are typically “v” shaped in cross 

section, and include features such as steep walls, 

exposed outcroppings of bedrock and clay, and 

tributaries. Most canyons may have been formed 

by mass-wasting processes on the continental 

slope; some, like the Hudson Canyon (MidAtlantic), may have formed because of fluvial 

drainage. The canyons exhibit a more diverse 

fauna, topography, and hydrography than the 

surrounding shelf and slope environments. The 

diversity in substrata types tends to make the 

canyons biologically richer than the adjacent 

shelf and slope.

As mentioned above, the New England Seamount 

chain extends southeast of Georges Bank for 

about 1,100 km, rising as much as 4,000 m above 

the Sohm Abyssal Plain. Of the four seamounts 

within the U.S. EEZ, Bear Seamount is the closest 

and rises from a depth of 2,000-3,000 m to a 

summit that is 1,100 m below the sea surface. 

The minimum depths of the others are: Physalia 

(1,848 m), Mytilus (2,269 m), and Retriever 

(1,819 m). Several other seamounts outside the 

U.S. EEZ are biologically significant because 

they rise to relatively shallow depths. Substrate 

types range from solid basalt to manganese 

crusts to rock and coral rubble to mixtures of 

basalt pebbles and sand to fine carbonate oozes. 

Sediments cover the summits with more exposed 

rock surfaces on the sides.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

Mid-Atlantic Bight

The Mid-Atlantic Bight refers to the region of the 

continental shelf and slope waters from Georges 

Bank south to Cape Hatteras, and east to the Gulf 

Stream. Here the shelf topography is relatively 

smooth, as depth increases linearly from shore 

to shelf break, except near submarine canyons. 

The primary features of the shelf include shelf 

valleys and channels, shoal massifs, scarps, 

and sand ridges. The main physiographic feature 

within the Mid-Atlantic is the Hudson Shelf 

Valley and Canyon, extending from the innercontinental shelf, at about the 40 m isobath, 

onto the continental slope. Other significant 

physiographic features include several other 

major canyons between Cape May and Cape 

Charles (Norfolk, Baltimore, Washington, and 

Wilmington), and the unique oceanography and 

geology off Cape Hatteras. Sediments over the 

Mid-Atlantic shelf are fairly uniformly distributed, 

and are primarily composed of sand, with 

isolated patches of coarse-grained gravel and 

fine-grained silt and mud deposits. Sand and 

gravel deposits vary in thickness from 0-10 m. 

The sands are mostly medium to coarse grains, 

with fine sand, silt, and clay in the Hudson shelf 

valley and on the outer shelf. Mud is rare over 

most of the shelf, but is common in the Hudson 

valley. Fine sediment content increases rapidly 

at the shelf break, which is sometimes called the 

“mud line,” and sediments are 70-100% fines 

on the slope. The continental slope off Cape 

Hatteras also receives exceptionally high fluxes 

of sediment and nutrients that are funneled off 

the shelf above, helping to account for the high 

abundance of infaunal organisms found there.

III. OCEANOGRAPHIC SETTING

Gulf of Maine

The Northeast Channel between Georges Bank 

and Browns Bank leads into Georges Basin, and 

is one of the primary avenues for water exchange 

between the Gulf of Maine and Atlantic Ocean. 

The Gulf has a general counterclockwise nontidal 

surface current that flows around the margin of 

the Gulf along the shore. This current is primarily 

driven by fresh, cold Scotian shelf water that 

enters from the north and through the Northeast 

Channel, and freshwater runoff from coastal 

rivers, which is particularly important in the 

spring. Dense, relatively warm and saline slope 

water entering through the Northeast Channel 

from the continental slope also influences gyre 

formation. Gulf circulation can vary significantly 

from year to year due to shelf-slope interactions 

such as the entrainment of shelf water by Gulf 

Stream rings, strong winds which can create fast 

moving currents, and annual and seasonal inflow 

variations. In the summer, the water in Jordan, 

Wilkinson, and Georges Basins becomes layered 

into warm, nutrient-poor surface water; cold, 

nutrient-rich intermediate water; and cool highsalinity bottom water.

Georges Bank

There is a persistent clockwise gyre around 

Georges Bank, a strong semidiurnal tidal flow 

predominantly northwest and southeast, and 

very strong, intermittent storm-induced currents, 

which can all occur simultaneously. Tidal currents 

over the shallow top of the Bank can be very 

strong, and keep the waters over the Bank well 

mixed vertically. This results in a tidal front that 

separates the cool waters of the well-mixed 

shallows from the warmer, seasonally stratified 

shelf waters on the seaward and shoreward sides 

of the Bank. The clockwise gyre is instrumental 

in distribution of the planktonic community, 

including larval fish. Georges Bank has a diverse 

biological community that is influenced by many 

environmental conditions and is characterized 

by high levels of primary productivity and, 

historically, high levels of fish production, which 

includes such species as Gadus morhua (Atlantic 

cod), Melanogrammus aeglefinus (haddock), and 

Limanda ferruginea (yellowtail flounder).

Mid-Atlantic Bight

The shelf and slope waters of the Mid-Atlantic 

Bight have a slow southwestward flow that is 

occasionally interrupted by warm core rings or 

meanders from the Gulf Stream. Slope water 

tends to be warmer and more saline than shelf 

water. The abrupt gradient where these two water 

masses meet is called the shelf-slope front. This 

front is usually located at the edge of the shelf 

and touches bottom at about 75-100 m depth of 

water, and then slopes up to the east toward the 

surface. It reaches surface waters approximately 

25-55 km further offshore. The position of the 

front is highly variable, and can be influenced by 

many physical factors.
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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a complete list of the deep coral species in this 

region, see Appendix 5.1.

a. Stony Corals (Class Anthozoa, Order 

Scleractinia)

Cairns and Chapman (2001) list 16 species 

of stony corals from the Gulf of Maine and 

Georges Bank to Cape Hatteras (Table 5.1) 

(See also Cairns 1981). Most of the stony corals 

in this region are solitary organisms and one 

species, Astrangia poculata, can occur in very 

shallow water, at depths of only a few meters. 

Moore et al. (2003, 2004) reported several 

species of solitary and colonial stony corals on 

Bear Seamount; one notable solitary species, 

IV. STRUCTURE AND HABITAT-FORMING 

DEEP CORALS

As stated in the introduction, the northeast region 

does not appear to have an abundance of large, 

structure-forming deep corals and deep coral 

habitats. Thus, we will confine our discussions 

primarily to the major deep gorgonian corals as well 

as to some of the more noteworthy occurrences 

of alcyonaceans (soft corals) and scleractinians 

(stony corals), including solitary species or 

those found in shallower habitats. Records of 

gorgonians and soft corals are often combined 

into one database, so for convenience we will be 

discussing those two orders simultaneously. For 

Table 5.1. Stony corals from the Gulf of Maine and Georges Bank to Cape Hatteras, based on Cairns and 

Chapman (2001). Vaughanella margaritata has been reported from Bear Seamount of the New England Seamount 

chain (Moore et al. 2003). Depth ranges are for the western Atlantic only. * = potentially structure-forming colonial 

species.

Taxon Distribution Coloniality Attachment Depth (m)

Astrangia poculata (Ellis & 

Solander, 1786) Endemic to western Atlantic colonial attached 0-263

Caryophyllia ambrosia 

ambrosia Alcock, 1898

Widespread (cosmopolitan) 

distribution solitary free-living 1487-2286

Caryophyllia ambrosia 

caribbeana Cairns, 1979 Endemic to western Atlantic solitary free-living 183-1646

Dasmosmilia lymani

(Pourtales, 1871)

Widespread (cosmopolitan) 

distribution solitary free-living 37-366

Deltocyathus italicus

(Michelotti, 1838)

Amphi-Atlantic with a 

disjunct distribution solitary free-living 403-2634

Desmophyllum dianthus 

(Esper, 1794)

Widespread (cosmopolitan) 

distribution solitary attached 183-2250

Enallopsammia profunda

(Pourtales, 1867)* Endemic to western Atlantic colonial attached 403-1748

Enallopsammia rostrata

(Pourtales, 1878)*

Widespread (cosmopolitan) 

distribution colonial attached 300-1646

Flabellum alabastrum

Moseley, 1873

Amphi-Atlantic with 

contiguous distribution solitary free-living 357-1977

Flabellum angulare Moseley, 

1876

Amphi-Atlantic with 

contiguous distribution solitary free-living 2266-3186

Flabellum macandrewi Gray, 

1849

Amphi-Atlantic with 

contiguous distribution solitary free-living 180-667

Fungiacyathus fragilis Sars, 

1872

Widespread (cosmopolitan) 

distribution solitary free-living 412-460

Javania cailleti (Duch. & 

Mich., 1864)

Widespread (cosmopolitan) 

distribution solitary attached 30-1809

Lophelia pertusa (L, 1758)* Widespread (cosmopolitan) 

distribution colonial attached 146-1200

Solenosmilia variabilis

Duncan, 1873*

Widespread (cosmopolitan) 

distribution colonial attached 220-1383

Vaughanella margaritata 

(Jourdan, 1895)

Endemic to northwestern 

Atlantic solitary attached 1267
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

Vaughanella margaritata, represents 

the first record of this species since its 

original description over 100 years ago, 

and is endemic to the northwest Atlantic 

(Cairns and Chapman 2001). Other 

recent expeditions to the New England 

and Corner Rise Seamounts have also 

found stony corals (Adkins et al. 2006; 

Watling et al. 2005; Shank et al. 2006).

b. Black Corals (Class Anthozoa, 

Order Antipatharia)

Antipatharians are predominantly 

tropical, but some species are known 

to occur in the northwest Atlantic. 

Bushy black coral (Leiopathes sp.) has 

been collected from 1643 m on Bear 

Seamount (Brugler 2005); it is also found 

in the collections of the Smithsonian 

Institution, having been collected in 

1883 by the R/V Albatross from 1754 m 

near the same area off Georges Bank. 

Within the New England Seamount 

chain, very few associated species 

have been found living on Leiopathes

sp. (Brugler 2005). Another black coral, 

Cirrhipathes sp., is also found in the 

Smithsonian Institution collections, and 

was also collected in 1883 by the R/V 

Albatross at 262 m off Virginia. Watling 

et al. (2005) collected at least 8 species 

of black coral from the seamounts during 

their 2004 expedition; Brugler and 

France (2006) observed and collected 

15 species of black coral during their 

2005 expedition to the New England 

and Corner Rise Seamounts, including 

7 species that they did not previously 

observe on the seamounts.

c. Gold Corals (Class Anthozoa, 

Order Zoanthidea)

No records of species from this order 

have been found in this region.

d. Gorgonians (Class Anthozoa, 

 Order Gorgonacea)

The Watling et al. (2003) database obtained 

records of both gorgonian and true soft 

coral occurrences from a variety of sources, 

including Verrill, Deichmann (1936), Hecker 

and collaborators [e.g.; Hecker (1980, 1990), 

Hecker and Blechschmidt (1980), Hecker et 

al. (1980, 1983); see descriptions below], Yale 

Peabody museum collections, the NEFSC 

benthic database of identified coral taxa, and 

observations from recent National Undersea 

Research Center (NURC) field studies [for further 

information, see Watling and Auster (2005)]. A 

total of 17 species in 7 families were recorded 

for the northeastern U.S. shelf and slope north of 

Cape Hatteras (Table 5.2). These 17 species in 

the seven gorgonian families (Acanthogorgiidae, 

Paramuriceidae, Anthothelidae, Paragorgiidae, 

Taxa Species

Order Gorgonacea

Acanthogorgiidae Acanthogorgia armata Verrill, 1878

Paramuriceidae Paramuricea grandis Verrill, 1883

Paramuricea placomus (Linné, 1758)

Paramuricea n. sp.

Swiftia casta (Verrill, 1883)

Anthothelidae Anthothela grandiflora (Sars, 1856)

Paragorgiidae Paragorgia arborea (Linné, 1758)

Chrysogorgiidae Chrysogorgia agassizii (Verrill, 1883)

Iridogorgia pourtalesii Verrill, 1883

Radicipes gracilis (Verrill, 1884)

Primnoidae Narella laxa Deichmann, 1936

Primnoa resedaeformis (Gunnerus, 1763)

Thouarella grasshoffi Cairns, 2006

Isididae Acanella arbuscula (Johnson, 1862)

Keratoisis ornata Verrill, 1878

Keratoisis grayi Wright, 1869

Lepidisis caryophyllia Verrill, 1883

Order Alcyonacea

Clavulariidae Clavularia modesta (Verrill, 1874)

Clavularia rudis (Verrill, 1922)

Alcyoniidae Alcyonium digitatum Linné, 1758

Anthomastus grandiflorus Verrill, 1878

Anthomastus agassizii Verrill, 1922

Nephtheidae Gersemia rubriformis (Ehrenberg, 1934)

Gersemia fructicosa (Sars, 1860)

Capnella florida (Rathke, 1806)

Capnella glomerata (Verrill, 1869

Table 5.2. List of gorgonians and soft corals known to occur 

on the northeastern U.S. continental shelf and slope north of 

Cape Hatteras, from Watling and Auster (2005), with taxonomic 

changes based on Integrated Taxonomic Information System 

(ITIS 2006) database/website and S. Cairns (Smithsonian Institution, Washington, D.C., pers. comm.).
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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Table 5.3. Identity and distribution of Pennatulacea on the northeastern U.S. continental shelf and slope. NMNH = Smithsonian Natural Museum of Natural History; OBIS = Ocean Biogeographic Information System.

Higher Taxon Species Distribution Depth Range (m) Reference

Anthoptilidae Anthoptilum grandiflorum Newfoundland to Bahamas, Lousiana, Chile, Hawaii, Antarctica, N. Europe 274-3651 U.S. NMNH collection, OBIS

Anthoptilum murrayi Lydonia Canyon to Puerto Rico, Hawaii, Aleutians, Japan, W. Africa, N. Europe 430-2491 (1538 m min in NE U.S.) U.S. NMNH collection, OBIS

Kophobelemnidae Kophobelemnon stelliferum Newfoundland to South Carolina, Japan, W. Africa, N. Europe 393-2199 (1330 m min in NE U.S.) U.S. NMNH collection, OBIS

Kophobelemnon scabrum Nova Scotia to Virginia 1977-2249 U.S. NMNH collectionKophobelemnon tenue Massachusetts to Virginia 2491-4332 U.S. NMNH collection

Pennatulidae Pennatula aculeata Newfoundland to Virginia, California, Iberia, N. Africa 119-3316 U.S. NMNH collection, OBIS

Pennatula grandis New Jersey, Bay of Biscay 1850-2140 U.S. NMNH collection, OBISPennatula borealis Newfoundland to North Carolina, California219-2295 U.S. NMNH collection, OBIS

Protoptilidae Distichoptilum gracile Nova Scotia to North Carolina, W. Africa, N. Europe 1211-2844 (doubtful report at 59 m) U.S. NMNH collection, OBIS

Protoptilum abberans Nova Scotia to Virginia 1483-2359 U.S. NMNH collectionProtoptilum carpenteri Massachusetts to North Carolina, W. Africa, N. Europe 1334-2194 U.S. NMNH collection, OBIS

Scleroptilidae Scleroptilum gracile Massachusetts to Virginia 2513-4332 U.S. NMNH collection

Scleroptilum grandiflorum Massachusetts to North Carolina, Panama, W. Africa 1502-2505 U.S. NMNH collection, OBIS

Umbellulidae Umbellula guntheri Massachusetts to Virginia, Louisiana 2683-3740 (3166 m min in NE U.S.) U.S. NMNH collection

Umbellula lindahlii Massachusetts to the Virgin Islands, Louisiana, Suriname, N. Europe, Indian O. 549-3338 (1538 m min in NE U.S.) U.S. NMNH collection, OBIS

Virgulariidae Balticina finmarchica Newfoundland to Massachusetts, NC (doubtful), the Virgin Islands, Alaska 37-2249 (229 m min in NE U.S.) U.S. NMNH collection

Stylatula elegans New York-Florida, Iberia 20-812 (51 m min in NE U.S.) U.S. NMNH collection, OBIS
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

Figure 5.2 A. Distribution of stony or hard 

corals (Astrangia, Dasmosmilia, and Flabellum) in the Gulf of Maine and Georges 

Bank. B) Distribution of stony or hard corals 

(Astrangia, Dasmosmilia, and Flabellum) in 

the Mid-Atlantic. Source credit: Theroux and 

Wigley (1998) database.

A

B

e. True Soft Corals (Class Anthozoa, 

 Order Alcyonacea)

As stated above, the Watling et al. (2003) 

database obtained records of both 

gorgonian and true soft coral occurrences 

from a variety of sources, including 

Verrill, Deichmann (1936),Hecker and 

collaborators [e.g.; Hecker (1980, 1990), 

Hecker and Blechschmidt (1980), Hecker 

et al. (1980, 1983); see descriptions 

below], Yale Peabody museum 

collections, the NEFSC benthic database 

of identified coral taxa, and observations 

from recent National Undersea Research 

Center (NURC) field studies [for further 

information, see Watling and Auster 

(2005)]. A total of 9 species in 3 families 

were recorded for the northeastern U.S. 

shelf and slope north of Cape Hatteras 

(Table 5.2). Two species that were not 

common in the database, but apparently 

are very numerous in nearshore records, 

were the soft corals Gersemia rubiformis 

and Alcyonium species (Watling and 

Auster 2005). As with the gorgonians, 

it should be noted that, for a variety 

Chrysogorgiidae, Primnoidae, and Isiddae) 

are the best documented because of 

their larger sizes, as well as being most 

abundant in the deeper waters of the 

continental slope (Watling and Auster 

2005). It should be noted that, for a variety 

of reasons, there is uncertainty about the 

accuracy of the identifications from these 

various surveys (Watling and Auster 2005), 

so these surveys should be interpreted 

with caution. Gorgonians have also been 

collected during recent expeditions to the 

New England and Corner Rise Seamounts 

(Moore et al. 2003, 2004; Watling et al. 

2005, 2006).
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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Figure 5.3 A) Distribution of gorgonians and 

soft corals in the Gulf of Maine and Georges 

Bank. B) Distribution of gorgonians and soft 

corals in the Mid-Atlantic. Source credits: Theroux and Wigley (1998) database, Watling et al. 

(2003) database.

A

B

of reasons, there is uncertainty about 

the accuracy of the identifications from 

these various surveys (Watling and 

Auster 2005), so these surveys should 

be interpreted with caution. Soft corals 

have also been collected during recent 

expeditions to the New England and 

Corner Rise Seamounts (Moore et al. 

2003; Watling et al. 2005, 2006).

f. Pennatulaceans (Class Anthozoa, 

Order Pennatulacea)

Records of sea pens were drawn from 

Smithsonian Institution collections and 

the Theroux and Wigley benthic database. 

Nearly all materials from the former 

source were collected either by the U.S. 

Fish Commission (1881-1887) or for the 

Bureau of Land Management (BLM) by 

the Virginia Institute of Marine Sciences 

(1975-1977) and Battelle (1983-1986). 

These latter collections heavily favor the 

continental slope fauna. The Theroux 

and Wigley collections (1955-1974) were 

made as part of a regional survey of all 

benthic species (Theroux and Wigley 

1998), heavily favoring the continental 

shelf fauna. A list of 17 sea pen species 

representing five families was compiled 

from these sources for the northeastern 

U.S. (Table 5.3). None commonly occur in 

shallow water, and only two species are 

known from the lower continental shelf 

depths (80-200 m) in this region: Pennatula 

aculeata (common sea pen) and Stylatula 

elegans (white sea pen). P. aculeata is 

common in the Gulf of Maine (Langton et 

al. 1990), and there are numerous records 

of Pennatula sp. on the outer continental 

shelf as far south as the Carolinas in the 

Theroux and Wigley database. S. elegans

is abundant on the Mid-Atlantic coast outer 

shelf (Theroux and Wigley 1998). The other 

15 sea pen species have been reported 

exclusively from the continental slope 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

Figure 5.4 A) Distribution of the gorgonians 

Acanthogorgia armata, Paragorgia arborea, and 

Primnoa resedaeformis in the Gulf of Maine and 

Georges Bank. B) Distribution of the gorgonians 

Acanthogorgia armata, Paragorgia arborea, 

and Primnoa resedaeformis in the Mid-Atlantic. 

Source credit: Watling et al. (2003) database.

A

B

depths (200-4300 m). Unlike most other 

corals, sea pens live in muddy sediments, 

anchored in place by a swollen, buried 

peduncle. Some species are capable of 

retracting part or the entire colony into the 

sediment when disturbed.

g. Stylasterids (Class Hydrozoa, Order 

Anthothecatae, Suborder Filifera, 

Family Stylasteridae)

No records of species from this family have 

been found in this region (Cairns 1992).

V. SPATIAL DISTRIBUTION 

OF CORAL SPECIES AND 

HABITATS

General Distribution in the Gulf of Maine, 

Georges Bank, and Mid-Atlantic Bight

Theroux and Wigley (1998) described the 

distribution of deep corals in the northwest 

Atlantic, based on samples taken from 

1956-1965. They often do not distinguish 

between taxonomic groups; e.g., stony 

corals such as Astrangia sp. and Flabellum

sp. are lumped together with the various 

types of anemones in the subclass 

Zoantharia. The distributions of only 

the stony corals, specifically Astrangia, 

Dasmosmilia, and Flabellum, from the 

Theroux and Wigley (1998) database in 

the Gulf of Maine/Georges Bank, and 

Mid-Atlantic are depicted in Figures 5.2A 

and B, respectively. There appears to be 

a general lack of stony corals on Georges 

Bank (Figure 5.2A), but note their presence 

along the continental margin (Figure 5.2A 

and B).
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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A

C

B

Figure 5.5 Distribution of the gorgonians Acanthogorgia armata, Paragorgia arborea, and Primnoa resedaeformis in A) Lydonia and B) Oceanographer Canyons (Gulf of Maine) and C) Baltimore 

Canyon (Mid-Atlantic). Source credit: Watling et al. 

(2003) database.

Theroux and Wigley (1998) also discussed the 

soft corals, gorgonians, as well as the sea pens. 

They were present along the outer margin of the 

continental shelf and on the slope and rise, and 

were sparse and patchy in all areas, particularly in 

the northern section. Theroux and Wigley (1998) 

found that they were not collected in samples 

taken at <50 m in depth, and were most abundant 

between 200-500 m. Figures 5.3A and B depict 

the distribution of gorgonians and soft corals in 

the Gulf of Maine, Georges Bank, and the MidAtlantic, based on the Theroux and Wigley (1998) 

database and the Watling et al. (2003) database. 

Identified species include gorgonians such as 

Acanella sp., Paragorgia arborea, Primnoa 

reseda [now resedaeformis, see Cairns and 

Bayer (2005)] and the soft coral Alcyonium sp. 

Gorgonians and soft corals were collected from 

gravel and rocky outcrops (Theroux and Wigley 

1998).

Watling and Auster (2005) noted two distinct 

distributional patterns for the gorgonians and 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

soft corals. Most are deepwater species that 

occur at depths >500 m; these include species 

of gorgoninans in the genera Acanthogorgia, 

Acanella, Anthothela, Lepidisis, Radicipes, 

and Swiftia, and soft corals in the genera

Anthomastus and Clavularia. Other species 

occur throughout shelf waters to the upper 

continental slope and include the gorgonians P. 

arborea, P. resedaeformis, and species in the 

genus Paramuricea. P. arborea was described 

by Wigley (1968) as a common component of the 

gravel fauna of the Gulf of Maine, while Theroux 

and Grosslein (1987) reported P. resedaeformis, 

as well as P. arborea, to be common on the 

Northeast Peak of Georges Bank. Both species 

are widespread in the North Atlantic (Tendal 

1992); P. resedaeformis has been reported south 

to off Virginia Beach, Virginia (37°03’N) (Heikoop 

et al. 2002).

Figures 5.4A and B depict the distribution of 

three major gorgonians: Acanthogorgia armata, 

P. arborea, and P. resedaeformis in the Gulf 

of Maine/Georges Bank and the Mid-Atlantic, 

respectively, based on the Watling et al. (2003) 

database. All three species occur in Lydonia and 

Oceanographer Canyons located off Georges 

Bank (Figure 5.4A), and Baltimore and Norfolk 

Canyons in the Mid-Atlantic (Figure 5.4B). Figure 

5.1 shows the locations of the major submarine 

canyons off the New England/Mid-Atlantic U.S. 

coast. The majority of records for these three 

species come from Lydonia, Oceanographer, 

and Baltimore canyons (Figure 5.5A, B, and 

C respectively). In addition, P. resedaeformis

was found throughout the Gulf of Maine and on 

the Northeast Peak of Georges Bank (Figure 

5.4A), affirming Theroux and Grosslein’s (1987) 

observations. 

It should be noted that the distribution maps 

presented in this chapter, based on both the 

Theroux and Wigley (1998) and Watling et al. 

(2003) databases, show presence only; i.e., 

they only describe where deep corals that could 

be identified were observed or collected. Since 

all areas have not been surveyed and since 

some specimens were not identified, the true 

distributions of many of these species remain 

unknown. However, the combination of these 

two databases represents the best available 

georeferenced data on the presence of deep 

corals in this region.

Continental Margin/Slope and Submarine 

Canyons

Dr. Barbara Hecker and her colleagues surveyed 

the deep corals and epibenthic fauna of the 

continental margin and several canyons off the 

northeastern U.S. in the 1980s via submersible 

and towed camera sled (Hecker et al. 1980, 

1983). Corals were denser and more diverse in 

the canyons, and some species, such as those 

restricted to hard substrates, were found only in 

canyons while the soft substrate types were found 

both in canyons and on the continental slope 

(Hecker and Blechschmidt 1980). The following 

is a summary of their findings for several of the 

prominent submarine canyons and continental 

slope areas; the surveys of other researchers are 

also included.

Lydonia Canyon

In the axis of Lydonia Canyon, deep corals were 

a major component of the fauna, and most were 

restricted to hard substrates (Hecker et al. 1980). 

The gorgonians Paramuricea grandis and A. 

armata were found in the axis and on the walls, with 

P. grandis being more common in the deeper part 

of the axis at depths >800 m. Other gorgonians

on hard substrates occurring along the axis of the 

canyon included P. arborea, P. resedaeformis, 

and Anthothela grandiflora, as well as the soft 

coral Trachythela (now Clavularia) rudis (Hecker 

et al. 1980). The most abundant coral found was 

the soft coral Eunephthya (now Capnella) florida, 

which was most common farther up the east 

axis between 500-700 m. Several individuals of 

a closely related species, Capnella glomerata, 

occurred on both the east and west walls of the 

canyon. The solitary stony coral Desmophyllum 

cristagalli (= dianthus) was found on outcrops in 

the deeper part of the axis and in some areas 

of the west wall. Other stony corals found in the 

canyon include Dasmosmilia lymani, Flabellum 

sp., Javania cailleti and Solenosmilia variabilis

(Hecker et al. 1983). The availability of suitable 

substrate appeared to be the most important 

factor in determining the distribution of the corals 

in Lydonia Canyon; for example, D. dianthus and 

P. grandis, which were common inhabitants of 

outcrops along the canyon axis, were also found 

on outcrops and boulders on the slope (Hecker 

et al. 1983).

Oceanographer Canyon

On the west flank of Oceanographer Canyon, 

glacial erratics and coral rubble provide hard 
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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substrate for the attachment of several species of 

coral, the most common being the gorgonian A. 

armata from 650-950 m (Hecker and Blechschmidt 

1980; Hecker et al. 1980). A. armata and P. 

grandis were also found on the east flank, and on 

both the east and west walls along the axis. The 

soft coral C. florida was also abundant on the east 

flank. The most common species in the deeper 

zone was the gorgonian P. grandis (Hecker et 

al. 1980). Hecker and Blechschmidt (1980) and 

Valentine et al. (1980) also noted the presence 

of the Paramuricea borealis (now P. grandis), in 

Oceanographer Canyon; Valentine et al. (1980) 

observed their greatest abundance to be from 

1100-1860 m, while Hecker and Blechschmidt 

(1980) observed that they were dominant from 

950-1350 m. A. armata was most common on 

smaller cobbles, boulders, and coral rubble while 

P. grandis was usually found on large boulders 

or outcrops (Hecker and Blechschmidt 1980; 

Hecker et al. 1980). Other deep corals on hard 

substrates in Oceanographer Canyon included 

the soft coral Anthomastus agassizii in the axis 

and on both walls, the soft coral Clavularia 

rudis on the west wall, large colonies of the 

gorgonian P. arborea in the axis above 1000 m, 

and numerous individuals of the small encrusting 

gorgonian, A. grandiflora, along the axis. 

Valentine et al. (1980) also found A. agassizii in 

a zone of greatest abundance from 1100-1860 

m, while Hecker and Blechschmidt (1980) found 

this species mostly from 950-1350 m on glacial 

erratics, outcrops, and coral rubble. The solitary 

stony coral, D. dianthus, was found throughout 

the axis between 1500-1600 m (Hecker and 

Blechschmidt 1980) and on the west flank. Deep 

corals restricted to soft substrates included the 

soft coral Anthomastus grandiflorus on the east 

flank (Hecker and Blechschmidt 1980; Hecker et 

al. 1980) and the gorgonian Acanella arbuscula

on both walls (Hecker and Blechschmidt 1980; 

Hecker et al. 1980); A. arbuscula was found by 

Hecker and Blechschmidt (1980) mostly from 

950-1350 m. The gorgonian P. resedaeformis is 

also found in the canyon, with a zone of greatest 

abundance from 300-1099 m (Valentine et al. 

1980).

Baltimore Canyon

Compared to Lydonia and Oceanographer 

Canyons, Baltimore Canyon in the MidAtlantic had the fewest corals, perhaps due to 

the scarcity of exposed outcrops (Hecker et al. 

1980). At depths >400 m, where the canyon axis 

constricts and bends, outcrops and talus blocks 

are exposed. Several corals restricted to hard 

substrates were found in this area by Hecker et 

al. (1980). Massive colonies of the gorgonian P. 

arborea were found on the large rock outcrops. 

Other corals found included the gorgonians 

A. armata, P. resedaeformis, A. grandiflora, A. 

arbuscula, and the soft corals C. florida and A. 

agassizii (Hecker et al. 1980, 1983). The solitary 

stony coral D. lymani occurred in dense localized 

patches near the head of Baltimore Canyon, but 

was absent from many other areas in the Canyon 

(Hecker et al. 1983). Other stony corals found 

included Flabellum sp. and D. dianthus (Hecker 

et al. 1983).

Other Canyons

Hecker and Blechschmidt (1980) surveyed the 

deep corals and epibenthic fauna of the several 

other canyons off the northeastern U.S. Discrete 

assemblages of corals were not identified. For a 

complete list of species found in the historical 

survey of Hecker and Blechschmidt (1980) 

and the Hecker et al. (1980) field study, see 

Appendix 5.2, which includes Opresko’s (1980) 

list of octocorals and Hecker’s (1980) list of 

scleractinians from those two surveys.

In Heezen Canyon, the gorgonian A. arbuscula

and the soft coral A. grandiflorus, both found 

on soft substrates, occurred at 850-1050 m; the 

gorgonian P. grandis was common from 1450-

1500 m; the soft coral A. agassizii and the stony 

coral D. dianthus were found from 1150-1500; D. 

dianthus was also found from 1500-1550 m. The 

walls of Corsair Canyon were heavily dominated 

by corals, all of which were restricted to soft 

substrates. The gorgonian A. arbuscula was 

prominent from 600-800 m, and the soft coral 

A. grandiflorus dominated from 800-1000 m. In 

Norfolk Canyon in the Mid-Atlantic, the stony coral 

D. dianthus and the gorgonian A. armata were 

found on hard substrate at 1050-1250 m; both 

were also observed in this canyon by Malahoff et 

al. (1982). Hecker and Blechschmidt (1980) also 

noted that the solitary stony coral Flabellum sp. 

was seen in high concentrations at 1300-1350 m 

depth in Norfolk Canyon, and the soft coral A. 

grandiflorus was found between 2150-2350 m.

Deep corals have been seen on the shelf around 

Hudson Canyon and in the head of the Canyon 

(see Appendix 5.2). For example, most recently 

a survey by Guida (NOAA Fisheries Service, 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

slope on the southwestern edge of Georges Bank, 

between Veatch and Hydrographer Canyons. 

In the Mid-Atlantic they surveyed two slope 

areas; one, called Slope Area I, was flanked by 

Lindenkohl Canyon on the south and Carteret 

Canyon on the north, and the other, called Slope 

II, was about 70 miles north of Slope I, and was 

bounded by Toms Canyon to the south and Meys 

Canyon to the north. Deep corals found on Slope 

III included the solitary stony corals D. lymani

and D. dianthus, the soft coral A. agassizii, and 

the gorgonian P. grandis. In the Mid-Atlantic, the 

solitary stony coral D. lymani occurred in very high 

abundances in both slope areas at depths <500 

m. Other stony corals found on Slope I included

Flabellum sp. and D. dianthus; those corals, as 

well as S. variabilis, were also found on Slope II. 

The gorgonians P. grandis and A. arbuscula and 

the soft coral A. agassizii were also present on 

both slopes.

NEFSC, James J. Howard Marine Sciences 

Laboratory, Highlands, NJ, unpublished data) 

of benthic habitats on the shelf around Hudson 

Canyon in 2001, 2002, and 2004 found the 

solitary stony coral D. lymani at a number of sites 

at depths of 100 to 200 m (Figure 5.6). They were 

particularly abundant in patches in a narrow band 

along the canyon’s rim near its head at depths of 

105-120 m; local densities within those patches 

exceeded 200 polyps m-2, but densities elsewhere 

were much lower. However, the only evidence of 

deep corals occurring deep within the canyon 

itself comes from Hecker and Blechschmidt 

(1980), who found abundant populations of 

the soft coral Eunephthya fruticosa (same as 

Gersemia fructicosa?) only in the deeper portion 

of the canyon.

Other Areas of the Continental Slope

Hecker et al. (1983) surveyed an area called 

Slope III, a 25 mile wide section of the continental 

Figure 5.6. Distribution and approximate densities (polyps per square meter) of the solitary stony coral 

Dasmosmilia lymani in samples from the Mid-Atlantic shelf around Hudson Canyon (Guida, unpublished data). 

Data obtained from still photos and trawl samples taken during October and November 2001, 2002, 2005, 

August 2004, and March 2007.
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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209

Hecker (1990), in a later survey of the megafaunal 

assemblages at four locations on the continental 

slope south of New England [including two that 

were surveyed as part of the Hecker et al. (1983) 

study] found that the solitary stony corals D. lymani

and Flabellum sp. dominated the fauna on the 

upper slope, although D. lymani was absent from 

their transect off Georges Bank. The gorgonian 

A. arbuscula and the soft coral C. florida, which 

dominated the fauna on the shallower section of 

the middle slope, were found only at their transect 

off Georges Bank; the soft coral A. agassizii was 

also found in dense populations there.

The New England Seamounts 

Deep corals are one of the dominant members 

of the epifaunal communities on the New 

England Seamount chain (Auster et al. 2005). 

A 2004 exploratory cruise to the New England 

Seamount chain revealed significant deep coral 

assemblages, with 27 octocoral species, 8 black 

coral species, and 2 stony coral species collected 

(Watling et al. 2005), including possible new 

species of gorgonians from the Chrysogorgiidae 

and Paragorgiidae families (Eckelbarger and 

Simpson 2005). A 2005 cruise to the New 

England and Corner Rise Seamounts sampled 

39 species of octocorals, including 7 that may be 

new to science, and observed and collected 15 

species of black coral, including 7 species that 

the researchers have not previously observed 

on the seamounts (Brugler and France 2006; 

Watling et al. 2006). Distributions of several 

species (e.g., Paragorgia sp., Lepidisis spp., 

Paramuricea spp., as well as stony corals and 

black corals) are currently being quantified using 

videotapes and digital still images (Figure 5.7). 

Preliminary quantitative analyses of coral species 

distributions indicate that community composition 

differs considerably between seamounts, 

even at comparable depths. These differences 

correspond to biogeographical boundaries, or 

they may be due to species’ responses to local 

habitat conditions, such as substratum type or 

flow. Substantial variation in faunal composition 

occurs between sites on a single seamount (P. 

Auster, pers. comm.).

During surveys of Bear Seamount within the 

EEZ during 2000, Moore et al. (2003) noted 

the presence of several species of stony 

corals, including the solitary corals Caryophyllia 

ambrosia, Flabellum alabastrum, and, as 

discussed previously, Vaughanella margaritata

and the colonial species Lophelia pertusa. 

Desmophyllum cristagalli (= dianthus) had been 

found on the Seamount during previous surveys. 

Gorgonians found by Moore et al. (2003) included 

Paragorgia sp., Lepidisis sp. Swiftia (?) sp., and 

Acanthogorgia angustiflora, although the latter 

species does not appear anywhere else in the 

deep coral records from this region. Although 

Moore et al. (2003) mentions that a species of 

Primnoa had been found on the Seamount during 

previous surveys, Cairns (Smithsonian Institution, 

Washington, D.C., pers. comm.) states that 

Primnoa does not occur on Bear Seamount; this 

record had originally been taken from Houghton 

et al. (1977), which is in error. Moore et al. (2003) 

also collected the soft coral A. agassizii.

Deep coral collected on Bear Seamount during 

a follow-up cruise in 2002 by Moore et al. (2004) 

included the solitary stony corals Caryophyllia 

cornuformis (this species is not mentioned 

elsewhere) and F. alabastrum and the colonial 

stony corals S. variabilis and Enallopsammia 

rostrata. Gorgonians collected included 

Paragorgia sp., Paramuricea sp., Keratoisis sp., 

Swiftia pallida (this species is not mentioned 

elsewhere), Lepidisis sp., and Radicipes gracilis.

VI. SPECIES ASSOCIATIONS WITH DEEP 

CORAL COMMUNITIES

Commercial Fisheries Species

The role of deep corals as possible habitats for 

fishes has only recently been addressed in the 

literature. The corals Primnoa, Lophelia, and 

Oculina from other regions have been the most 

studied. Several studies have documented that 

certain fish commonly occur in the vicinity of 

corals more often than in areas without corals. 

In the northwest Atlantic, this has been noted 

for Sebastes sp. (redfish) in the Northeast 

Channel (Mortensen et al. 2005). Redfish may 

take advantage of structure on the bottom as a 

refuge from predation, as a focal point for prey, 

and for other uses. However, in a survey of 

habitats in the Jordan Basin in the Gulf of Maine 

containing coral assemblages (primarily from the 

genera Paragorgia, Paramuricea, and Primnoa), 

Auster (2005) found that densities of redfish 

were not significantly different between dense 

coral habitats and dense epifauna habitats, 

although the density of redfish in these two 

habitats was higher than in the outcrop-boulder 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

habitat containing sparse epifauna. While this 

study shows that a habitat without deep corals 

can support similar densities of fish to a habitat 

containing corals, Auster (2005) states that 

it is the actual distribution of each habitat type 

throughout a region that will ultimately determine 

the role such habitats play in the demography of 

particular species and communities. Deep coral 

habitats are fairly rare in the Gulf of Maine (Figs. 

5.2A and 5.3A; Watling et al. 2003), but bouldercobble habitats containing dense epifauna are 

not. Auster (2005) suggests that deep corals 

do have some effect on the distribution and 

abundance of fishes, but by themselves do not 

support high density, unique or high diversity fish 

communities. The corals do provide important 

structural attributes of habitat, but may not be 

functionally different than structures provided by 

other dense epifaunal assemblages.

In addition, variations in the morphological forms 

of the deep corals themselves within a landscape 

can affect how they will be used by fishes. For 

example, on the New England Seamounts, Auster 

et al. (2005) describes the gorgonian assemblage 

deep on the seamount flanks as a mixture of 

“tall and whip-like species and short fans with 

low density branching,” with the taller species 

forming moderately dense stands, while near 

and on the seamount summits, the gorgonians 

(e.g., Paragorgia sp.) are often “large (~1 m 

wide) robust fans with high density branching” 

that are spread across the landscape. The form 

and density of the latter may have a different 

habitat value in terms of shelter or refuge from 

flow (Auster et al. 2005). Auster et al. (2005) did 

observe one fish species, Neocyttus helgae, that 

appeared to be associated with Paragorgia sp., 

perhaps utilizing them as a refuge from flow and 

as a foraging spot for prey associated with the 

coral, or perhaps they were foraging on the coral 

polyps themselves.

Invertebrates

There are few data available about invertebrate 

species associations with deep corals in 

this region; more is known about the species 

associations of deep corals and invertebrates 

from other regions [e.g., Primnoa off Alaska 

(Krieger and Wing 2002); Lophelia pertusa off 

Norway (Mortensen 2000)]. Off the northeastern 

U.S., Hecker et al. (1980) noted the frequent 

occurrence of shrimp on the largest gorgonians

such as Paragorgia arborea, Paramuricea 

grandis, Primnoa resedaeformis, and the soft 

coral Capnella florida. An ophiuroid and an 

Figure 5.7. Photographic examples of variations 

in coral communities from the upper slope of Bear 

Seamount, near the summit. A) Bamboo coral 

(Keratoisis sp.) at left with globular sponge (bottom center), brisingid sea star (center top) and 

leafy sponge (center right). B) Keratoisis sp. C) 

Lepidisis sp. Photo credit: Deep Atlantic Stepping 

Stones Science Team, IFE, URI, and NOAA-OE

A B

C
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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211

anemone appeared to have an association, 

possibly obligatory, as suggested by Hecker et 

al. (1980), with Paramuricea grandis. Hecker 

and Blechschmidt (1980) also noted several 

faunal associations in their study. The ophiuroids 

Ophiomusium lymani and Asteronyx loveni were 

often associated with corals; e.g., the latter with 

the gorgonian A. arbuscula. Pycnogonids were 

seen on the gorgonian Paramuricea grandis.

Current studies are looking at the species 

associations of the octocorals of the New 

England and Corner Rise Seamounts. Ophiuroids 

and marine scale worms were found to live 

commensally in and on the octocorals found on 

the seamounts (Watling 2005; Watling and Mosher 

2006); specimens of octocorals, stony corals, 

and their associated species from the seamounts 

are also being used in population genetic studies 

that will assess, for example, the taxonomic 

relationships of the corals and associated species 

(Shank et al. 2006). Within the EEZ, Moore et al. 

(2003) noted the following invertebrates found in 

direct association with Lophelia petusa trawled 

from Bear Seamount: a large polychaete worm 

living in tubular spaces within the coral colony, 

the attached solitary stony coral, Caryophyllia 

ambrosia, small serpulid worm tubes, a thecate 

hydroid, and the gorgonian Swiftia.

VII. STRESSORS ON DEEP CORAL 

COMMUNITIES

Fishing Effects

Deep corals provide habitat for other marine 

life, increase habitat complexity, and contribute 

to marine biodiversity, and their destruction 

could have a significant impact on other marine 

species. Anecdotal data from submersible and 

ROV studies as well as reports from fisherman, 

who have brought them up as bycatch since the 

19th century, suggests that deep corals have 

become less common or their distributions have 

been reduced due to the impacts of bottom 

fishing (Breeze et al. 1997; Watling and Auster 

2005). Fishing has had significant effects on deep 

coral populations in other regions. Deep corals 

are especially susceptible to damage by fishing 

gear because of their often fragile, complex, 

branching form of growth above the bottom. Also, 

they grow and reproduce at very slow rates, with 

some estimated to be hundreds of years old 

(Lazier et al. 1999; Andrews et al. 2002; Risk et 

al. 2002), and recruitment rates may also be low 

(Krieger 2001), which makes their recovery from 

disturbances difficult over short time periods. Of 

the various fishing methods, bottom trawling has 

been found to be particularly destructive (Rogers 

1999; Hall-Spencer et al. 2001; Koslow et al. 

2001; Krieger 2001; Fosså et al. 2002; Freiwald 

2002).

The effects of current and historic fishing 

efforts on deep coral and coral habitats in the 

northeastern U.S. have not been quantified. The 

types of fishing gear used here include fixed gear 

such as longlines, gillnets, and pots and traps, 

as well as trawls and dredges. Fixed gear can be 

lost at sea, where they can continue to damage 

corals. In Canada, longlines have been observed 

entangled in deep corals such as Paragorgia and 

Primnoa and may cause breakage (Breeze et 

al. 1997; Mortensen et al. 2005). Further, “ghost 

fishing” of nets entangled in coral reefs has been 

reported in the North Sea (ICES 2003). Bottom 

trawling was found to have a larger impact on 

deep corals per fishing unit of effort compared 

to longlining; e.g., damage to Primnoa off Alaska 

(Krieger 2001). The northeastern U.S. fisheries 

that have the highest likelihood of occurring near 

concentrations of known deep coral habitats 

(e.g., in canyon and slope areas) are the 

Lophius americanus (monkfish or goosefish) and 

Lopholatilus chamaeleonticeps (tilefish) fisheries, 

and the Chaceon (Geryon) quinquedens (red 

crab) and offshore Homarus americanus (lobster) 

pot fisheries.

Effects of other human activities

Invasive Species

Little is known about any existing or potential 

interactions between deep corals and invasive 

species. One invasive species that may be a 

threat in the northeast is the colonial tunicate (“sea 

squirt”), Didemnum sp. A. (Bullard et al. 2007). 

It is currently undergoing a massive population 

explosion and major range expansion and has 

become a dominant member of many subtidal 

communities on both coasts of North America. 

It was recently discovered on the northern edge 

of Georges Bank, and is found on hard gravel 

substrates, including, pebbles, cobbles, and 

boulders, and overgrows sessile and quasimobile epifauna (sponges, anemones, scallops, 

etc.). The species can occur on immobile sand 

substrates, but is not known to occur on mobile 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

sand or mud substrate. On Georges Bank, 

Didemnum sp. A covers 50–90% of the substrate 

at numerous sites over a 230 km2 area to a depth 

of 45-60 m (Bullard et al. 2007). Didemnum sp. 

A has the potential to spread rapidly by budding, 

and fragmentation of the mats could promote the 

rapid spread of the species. There are no known 

predators at this time, although there have been 

some observations of possible seastar predation 

(Bullard et al. 2007).

Didemnum sp. A may be a potential serious threat 

to deep coral that occur on hard substrates in the 

northeast, particularly for those corals, such as 

Paragorgia and Primnoa, which are known to 

occur on the gravel substrate of the Northeast 

Peak of Georges Bank. If the tunicate did 

colonize deep coral habitat, it could smother and 

kill the corals by overgrowth, or at least impede 

feeding (G. Lambert, University of Washington, 

Friday Harbor Laboratories, Seattle, WA, pers. 

comm.; P. Valentine, USGS, Woods Hole, MA, 

pers. comm.) However, Bullard et al. (2007) note 

that on cobble bottoms in Long Island Sound, 

the only epifaunal species that Didemnum sp. A 

did not appear to overgrow were the stony coral, 

Astrangia poculata, and the cerianthid anemone, 

Ceriantheopsis americana. In addition, depth, 

temperature, and larval dispersal may be limiting 

factors in the spread of the tunicate (G. Lambert, 

pers. comm.; P. Valentine, pers. comm.). It may be 

a shallow water species; so far, it has been found 

only at depths down to 81 m off New England on 

Georges, Stellwagen and Tillies Banks (Bullard 

et al. 2007), and even though it is found in a 

temperature range of -2°C to 24°C, its preferred 

range might be at temperatures >4°C or higher 

(G. Lambert, pers. comm.; P. Valentine, pers. 

comm.). Didemnum sp. A would probably have 

a difficult time getting from the present affected 

area into the submarine canyons located on the 

southern bank margin. The tadpole larvae that 

are born from sexual reproduction swim near the 

bottom (probably for only a few hours) before 

settling, so they cannot travel far on their own 

or by currents. However, because fragments 

of the colonies are viable, it could, in theory, 

be brought to the canyons on a boat hull or by 

the use of contaminated fishing gear (mobile or 

fixed) or the washing of contaminated boat decks 

(P. Valentine, pers. comm.). Even though the 

consensus is that Didemnum sp. A is probably 

a remote threat to deep corals, it has previously 

confounded researchers with its incredible rate 

of colonization and range expansion. So little is 

known about this tunicate that it is not possible to 

predict the limits of its further spread at this time 

(G. Lambert, pers. comm.). But Didemnum sp. A 

does has the potential to cause great ecological 

and economic damage. The species continues to 

expand its range and could eventually colonize 

large expanses of hard substrata throughout 

New England and eastern Canada (Bullard et al. 

2007).

Other

There does not appear to be any specific current 

or pending oil and gas exploration/extraction, gas 

pipeline/communication cable, mineral mining, 

etc. projects that could pose a significant threat 

to the major deep coral species in this region; 

in addition, few of the coastal projects along the 

northeast coast of the U.S. are deep enough to 

affect them. There also doesn’t appear to be any 

monitoring data or published studies from these 

types of projects in this region that show evidence 

of impacts to deep corals. 

VIII. MANAGEMENT OF FISHERY 

RESOURCES AND HABITATS

Mapping and Research

Despite the aforementioned faunal surveys, 

our knowledge of the temporal and spatial 

distribution and abundance of deep corals off the 

northeastern U.S., as well as some aspects of 

their basic biology and habitat requirements, is 

severely limited, so their overall population status 

and trends are difficult to determine. [There 

is, however, more information on deep coral 

distribution and habitat requirements in Canadian 

waters; e.g., the Northeast Channel (Mortensen 

and Buhl-Mortensen 2004)]. NEFSC groundfish 

and shellfish surveys from the Gulf of Maine to 

Cape Hatteras have collected corals as part of 

their bycatch for several decades, but there are 

many data gaps (e.g., corals were not properly 

identified or quantified) which prevent using the 

data to clearly assess any long-term population 

trends.

There have been some recent, targeted surveys 

off of New England using trawls and remotely 

operated vehicles (ROVs). In 2003, 2004, and 

2005, their were surveys of several seamounts 

in the New England and Corner Rise Seamount 

chains (the latter is approximately 400 km to 
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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the east of the New England Seamount chain, 

and nearly midway between the east coast of 

the U.S. and the Mid-Atlantic Ridge) funded by 

NOAA’s Office of Ocean Exploration and National 

Undersea Research Program. The cruises were 

multidisciplinary in nature but the goals included 

studying the distribution and abundance of deep 

corals relative to the prevailing direction of currents; 

collecting specimens for studies of reproductive 

biology, genetics, and ecology; and studying 

species associations. Mike Vecchione (NEFSC, 

National Systematics Laboratory) conducted 

a multi-year study (2000-2005) exploring the 

faunal biodiversity of Bear Seamount. The multiyear program examined some of the temporal 

variability around the Seamount. Initial results 

on the biodiversity of deep corals by Moore et al. 

(2003) were discussed previously. Mike Fogarty 

(NEFSC) in spring 2004 explored the macrofaunal 

biodiversity of the upper continental slope south 

of Georges Bank from Oceanographer Canyon 

to Powell Canyon at depths from 400-1100 m, 

with the deepest stations corresponding to the 

shallowest depths sampled on the summit of 

Bear Seamount in the Vecchione survey.

Fishery Management Councils

Fishery management council jurisdiction in the 

northeast U.S. is primarily the responsibility 

of the New England Fishery Management 

Council (NEFMC). In addition to the 26 species 

under its sole management, the NEFMC 

shares responsibility over Lophius americanus

(monkfish or goosefish) and Squalus acanthias

(spiny dogfish) with the Mid-Atlantic Fishery 

Management Council (MAFMC). In 2005, the two 

Councils, with the NEFMC as the lead Council on 

the Monkfish Fishery Management Plan (FMP), 

approved the designation of Oceanographer and 

Lydonia Canyons (approximately 116 square 

nautical miles) as Habitat Closed Areas (HCA) 

and added these areas to the NEFMC’s network 

of HCAs (or marine protected areas). These 

new HCAs are closed indefinitely to fishing with 

bottom trawls and bottom gillnets while on a 

monkfish day-at sea (DAS) in order to minimize 

the impacts of the directed monkfish fishery on 

Essential Fish Habitat (EFH1) in these deep-sea 

canyons and on the structure-forming organisms 

therein (Figures 5.1 and 5.8). Within these canyon 

habitats, a variety of species have been found 

which are known to provide structured habitat, 

including deep corals, and shelter for many 

species of demersal fish and invertebrates.

This action was implemented in May 2005 

under Amendment 2 to the Monkfish Fishery 

Management Plan (FMP) (NEFMC 2004). EFH 

for some federally-managed species extends 

beyond the edge of the continental shelf and 

includes portions of the canyons. 

The directed monkfish fishery is conducted with 

bottom trawls and bottom gillnets, primarily in 

coastal and offshore waters of the Gulf of Maine, 

on the northern edge of Georges Bank, and in 

coastal and continental shelf waters of southern 

New England, including offshore waters on the 

edge of the continental shelf and near the heads 

of several of the canyons. Although the current 

degree of overlap between the current monkfish 

fishing effort and the known presence of corals 

within the canyons is very small, one of the 

fishery management measures contained within 

Amendment 2, and which was approved by the 

Councils, would increase the probability that the 

offshore fishery for monkfish will expand in the 

future. Because there is documented evidence 

of deep corals in the canyons in the area that is 

identified for possible increased offshore fishing, 

these closures are intended as a precautionary 

measure to prevent any potential direct or indirect 

impacts of an expanded offshore monkfish fishery 

on EFH, offshore canyon habitats, and thus, deep 

corals.

Approximately 23 federally-managed species 

have been observed or collected within these 

proposed closure areas, and many of them have 

EFH defined as “hard substrates” at depths >200 

m, which includes habitat or structure-forming 

organisms such as deep corals. Also, the EFH 

designations for juvenile and adult life stages of 

six of these managed species (Sebastes spp., 

redfish, is one of them) overlap with the two 

1 EFH is a provision of the Magnuson-Stevens Fishery 

Conservation and Management Act (1996). The EFH 

provision states: “One of the greatest long-term threats 

to the viability of commercial and recreational fisheries 

is the continuing loss of marine, estuarine, and other 

aquatic habitats. Habitat considerations should receive 

increased attention for the conservation and management of fishery resources of the United States.” 

The definition of EFH in the legislation covers: “those 

waters and substrate necessary to fish for spawning, 

breeding, feeding, or growth to maturity.” The legislation mandates that NOAA Fisheries and the Councils 

implement a process for conserving and protecting 

EFH.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

Figure 5.8. The Oceanographer and Lydonia Canyon Essential Fish Habitat closed areas, under Amendment 

2 to the Monkfish Fishery Management Plan; locations of known alcyonaceans from the Theroux and Wigley 

(1998) and Watling et al. (2003) databases; and, 1999 and 2001 directed monkfish otter trawl trips. Source 

credit: 1999 and 2001 VTR).
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

NORTHEAST

215

closed areas, and EFH for all six of these species 

has been determined to be vulnerable to bottom 

trawling and perhaps also vulnerable to bottom 

gillnets. Although deep corals are not explicitly 

included in the EFH descriptions for any species 

in the northeast region, some deep corals are, 

of course, known to grow on hard substrates, 

and may themselves be considered a form of 

substrate. The rationale is that, since there are 

corals found within these proposed closed areas, 

this is indicative of areas of hard bottom. Also some 

coral species may provide the structural attributes 

of habitat similar to that provided by other dense 

epifaunal assemblages (as discussed above), 

and may be particularly vulnerable to damage 

or loss by trawling or gillnets. Thus, by avoiding 

any direct adverse impacts of bottom trawls and 

gillnets used in the monkfish fishery on EFH for 

the six species of fish and any indirect adverse 

impacts on hard bottom habitat and emergent 

epifauna, such as the deep corals, that grow in 

these habitats within the closed areas, adverse 

impacts of an expanded offshore fishery would 

be minimized to the extent practicable.

Protection of deep corals is a relatively new 

concept in this region and the NEFMC believes 

that there are several statutory and regulatory 

authorities that support the Councils’ initiative to 

protect deep coral habitats. The NEFMC took this 

proactive and precautionary approach to protect 

these sensitive habitats through aggressive 

fishery management measures, which is based 

on sound ecological principles, as appropriate 

and necessary. In Amendment 2 to the Monkfish 

FMP, the Councils, for the same reasons and 

rationale discussed above, also considered a 

management alternative that would have closed 

all 12 steep-walled canyons along the continental 

shelf-break from the Hague Line in the north to 

the North Carolina/South Carolina border in the 

south. Although this management alternative 

was not ultimately chosen by the Councils 

for implementation due to the lack of readily 

available coral data and potential negative social 

and economic impacts, the Councils did feel that 

the science and data supported the closure of 

Oceanographer and Lydonia Canyons at that 

time as a precautionary measure. The Councils 

determined that protection of deep corals in all 12 

canyons would be less certain than in just closing 

Lydonia and Oceanographer Canyons, until 

such time as additional surveys are conducted 

or evidence is examined which more thoroughly 

document the presence of corals in the other 10 

canyons. 

The New England Fishery Management Council 

alone has also indefinitely closed an additional 

3,000+ square nautical miles, as Habitat Closed 

Areas, in the Gulf of Maine, Georges Bank, and 

southern New England to bottom-tending mobile 

fishing gear to protect EFH (Figure 5.1), which 

indirectly protects any deep corals in those 

areas. This initial suite of HCAs was created 

under both Amendment 10 to the Atlantic Sea 

Scallop Fishery Management Plan in 2003 and 

Amendment 13 to the Northeast Multispecies 

Fishery Management Plan in 2004 as “Level 3” 

closures, and are closed indefinitely to all bottomtending mobile gear to protect EFH. The canyon 

habitat closures implemented under Amendment 

2 to the Monkfish FMP should be viewed as an 

addition to the suite of HCAs, and in concert 

with the other HCAs, provides a large network of 

marine protected areas (MPAs) as compared to 

the relatively small size of the geographic region 

under management.

Lastly, under the current effort to update all of the 

EFH provisions of all of the NEFMC FMPs, the 

NEFMC has approved a set of new Habitat Areas 

of Particular Concern (HAPCs2) including many 

steep-walled canyons on the eastern seaboard 

extending from the Hague Line to the North 

Carolina/South Carolina border and Bear and 

Retriever Seamounts to protect sensitive EFH 

and the habitat and structure-forming organisms 

therein. These new HAPCs will need approval 

from the MAFMC before they can be implemented 

through final action sometime in late 2008. The 

MAFMC will likely take up the topic at one of their 

upcoming meetings

IX. REGIONAL PRIORITIES TO 

UNDERSTAND AND CONSERVE DEEP 

CORAL COMMUNITIES

Mapping

Deep corals have been largely unmapped off the 

northeastern U.S, particularly in the Mid-Atlantic. 

What is currently known about coral distribution 

2

HAPCs are a subset of the much larger area identified 

as EFH that play a particularly important ecological 

role in the life cycle of a managed species or that are 

rare and/or particularly sensitive or vulnerable to 

human-induced environmental degradation and 

development activities.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

in this region is largely based on blind, random, 

or grid sampling with trawl gear, grab samplers, 

and drop cameras that was done twenty or more 

years ago. While the breadth of such surveys 

was vast, in most cases the density of data 

they generated is much too diffuse spatially and 

temporally to provide distributional data adequate 

for management purposes. Therefore, it is critical 

to identify, map, and characterize deep corals and 

their habitats, particularly in the canyons, utilizing 

more advanced technologies such as side-scan 

and multibeam sonars, manned submersibles and 

ROVs, towed camera sleds, etc. Low-resolution 

maps should be produced that cover large areas 

for purposes of identifying potential locations of 

deep corals, and high-resolution maps should 

be produced for site-specific areas where deep 

corals are known to exist (McDonough and 

Puglise 2003). Temporal/spatial changes in deep 

coral distribution and abundance need to be 

assessed, and long-term monitoring programs 

should be established.

Recently, Leverette and Metaxas (2005) 

developed predictive models to determine areas 

of suitable habitat for Paragorgia arborea and 

Primnoa resedaeformis along the Canadian 

Atlantic continental shelf and shelf break. 

Several environmental factors including slope, 

temperature, chlorophyll a, current speed and 

substrate were included in the analysis. Their 

results showed that the habitat requirements 

differed between the two gorgonians. P. arborea

occurred predominantly in steeply sloped 

environments and on rocky substrates, while the 

habitat for P. resedaeformis was more broadly 

distributed and located in areas with high current 

speed, rocky substrates and a temperature range 

between 5-10°C. The use of predictive modeling 

to generate habitat suitability maps and to identify 

suitable habitat for deep coral in the northeastern 

U.S. would be an important step toward deep 

coral conservation.

Research

It was stated previously that there is uncertainty 

about the accuracy of the identifications of 

deep corals from the various historical surveys. 

Identifying deep corals is difficult, and their 

taxonomy is often in question, so as a first step, 

some basic taxonomic issues need to be worked 

out. Molecular genetics is one tool that could be 

used, and this line of research may provide insight 

into coral larval dispersal. Genetic studies may 

also be useful for comparing corals regionally, 

nationally, and on either side of the Atlantic. 

For example, DNA-sequencing technology is 

currently being used to determine whether the 

corals around the New England Seamounts are 

endemic, or simply populations of species with 

broader geographic distributions; e.g., whether 

the corals are dispersing from the New England 

Seamounts into the deep Gulf of Maine and 

submarine canyons off Georges Bank. However, 

it’s important to note that there is a shortage of 

qualified coral taxonomists available to properly 

identify deep corals. With so few professional 

coral taxonomists, it will be difficult to make 

progress in deep coral mapping and distribution, 

for example. More students need to be trained in 

coral taxonomy at the graduate level, and more 

funding needs to be available for taxonomic 

research and to hire coral taxonomists at 

museums and universities.

In addition to taxonomy, basic life history studies 

on deep corals are needed in this region. There 

are fundamental questions on deep coral 

growth, physiology, reproduction, recruitment, 

recolonization rates, and feeding. Their habitat 

requirements need to be characterized. In 

addition, it is important to collect associated 

oceanographic, geologic, and other habitat 

parameter data in order to understand the 

physical parameters that affect the distribution 

and extent of deep coral habitats. Deep coral 

habitat biodiversity should be assessed, food web 

relationships need to be defined, and the role that 

the corals play in the life histories of associated 

species should be described and quantified. In 

terms of the latter, the possible role of deep corals 

as EFH for Federally managed species has to be 

determined. Finally, it is necessary to quantify 

the vulnerability or resilience of deep corals to 

various anthropogenic threats, especially from 

fishing, and to quantify the recovery rates of 

corals and coral habitats that have been injured 

or destroyed. Many of these recommendations 

for research on deep corals can be found in 

McDonough and Puglise (2003) and Puglise and 

Brock (2003).

The NEFSC needs to become more quantitative 

about their deep coral bycatch in the groundfish 

and shellfish surveys and fisheries observer 

program logs. Prior to the year 2000, for example, 

bycatch quantity in the NEFSC Placopecten 

magellanicus (Atlantic sea scallop) surveys 
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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217

were estimated by cursory visual inspection 

or “eyeballing” only (D. Hart, NOAA Fisheries 

Service, NEFSC, Woods Hole Laboratory, Woods 

Hole, MA, pers. comm.). The bycatch data for 

those surveys were divided up into 3 categories: 

substrate, shell, and other invertebrates; and the 

log sheets only recorded percent composition and 

total volume (bushels). In the fisheries observer 

program, the observers also log the presence 

of coral bycatch; however, they are lumped into 

one category (“corals and sponges”), and are 

not identified further. In addition, because the 

observer program observes thousands of trips 

every year in dozens of different fisheries, with 

each fishery having its own regulations for mesh 

size and configuration, a reported absence of 

coral at a location may simply be a function of the 

catchability of the gear used (D. Potter, Fisheries 

Sampling Branch, NOAA Fisheries Service, 

NEFSC, Woods Hole Laboratory, Woods Hole, 

MA, pers. comm.). This is also a problem with the 

NEFSC surveys; it is important to remember that 

fishing gear is not designed to “catch” corals. But 

at the very least, if there was an attempt made 

to properly identify coral bycatch from these two 

programs, one might come up with some form of 

presence data.

X. CONCLUSION

The overall quantity of deep coral habitat in the 

northeast region is unknown, and no systematic 

assessment of the distribution, abundance, and 

population dynamics of deep coral is available 

for this region. That, along with a dearth of 

information on their natural history, as well 

difficulties with their taxonomy, makes it difficult, 

if not impossible, to determine if there have been 

changes in deep coral occurrence or abundance 

over time. Nevertheless, even though there 

is no quantitative information on the extent of 

anthropogenic impacts to deep corals in this 

region, some of the areas where structure-forming 

deep corals are definitely known to occur (e.g., 

unique areas such as the submarine canyons 

and the New England Seamounts) are currently 

protected or under consideration for protection 

from bottom-tending fishing gear as EFH and 

Habitat Areas of Particular Concern.

Obviously, in order to better preserve and 

protect deep corals and deep coral habitat off 

the northeastern U.S., there needs to be 1) 

an increased mapping and survey effort at the 

Federal and academic level (including joint 

studies with Canada); 2) more basic research 

on deep coral taxonomy, life history, habitat 

requirements, species associations, etc.; and 

finally, 3) quantification on the susceptibility 

of deep corals to anthropogenic influences, 

particularly fishing.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

XI. REFERENCES

Adkins JF, Scheirer DP, Robinson L, Shank T, 

Waller R (2006) Habitat mapping of 

deep-Sea corals in the New England 

Seamount: populations in space and time. 

EOS Transactions, American Geophysical 

Union 87(36):suppl

Andrews AH, Cordes EE, Mahoney MM, Munk 

K, Coale KH, Cailliet GM, Heifetz J 

(2002) Age, growth, and radiometric age 

validation of a deep-sea, habitat forming 

gorgonian (Primnoa resedaeformis) from 

the Gulf of Alaska. In: Watling L, Risk 

M (eds.) Biology of cold water corals. 

Hydrobiologia 471:101-110

Auster P (2005) Are deep-water corals important 

habitats for fishes? Pages 643-656 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin, Heidelberg

Auster P, Moore J, Heinonen KB, Watling L 

(2005) A habitat classification scheme 

for seamount landscapes: assessing the 

functional role of deep-water corals as 

fish habitat. Pages 657-666 in Freiwald 

A, Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin, 

Heidelberg

Babb I (2005) Mountains in the sea: mission 

plan. Ocean Explorer/Explorations. 

National Oceanic and Atmospheric 

Administration. <http://oceanexplorer.

noaa.gov/explorations/03mountains/

background/plan/plan.html> 

Backus RH, Bourne DW, eds. (1987) Georges 

Bank. Cambridge, MA, USA: MIT Press. 

593 pp

Beardsley RC, Butman B, Geyer WR, Smith P 

(1997) Physical oceanography of the 

Gulf of Maine: an update. In: Wallace GT, 

Braasch EF (eds.) Proceedings of the Gulf 

of Maine Ecosystem Dynamics Scientific 

Symposium and Workshop. Regional 

Association for Research on the Gulf of 

Maine (RARGOM) Report 97-1:39-52

Breeze H, Davis DS, Butler M, Kostylev V (1997) 

Distribution and status of deep sea corals 

off Nova Scotia. Marine Issues Committee 

Special Publication Number 1, Ecology 

Action Center, Halifax, NS. 58 pp

Brooks DA (1996) Physical oceanography of the 

shelf and slope seas from Cape Hatteras 

to Georges Bank: a brief overview. Pages 

47-75. In: Sherman K, Jaworski NA, 

Smayda TJ (eds.) The Northeast Shelf 

Ecosystem -- assessment, sustainability, 

and management. Cambridge, MA, USA: 

Blackwell Science

Brugler M (2005) Linnaean terminology with a 

twist! Ocean Explorer/Explorations/North 

Atlantic Stepping Stones/August 14 

Log. National Oceanic and Atmospheric 

Administration. <http://oceanexplorer.

noaa.gov/explorations/05stepstones/

logs/aug14/aug14.html> 

Brugler M, France SC (2006) Distribution and 

abundance of black corals. Ocean 

Explorer/Explorations/North Atlantic 

Stepping Stones/Log. National Oceanic 

and Atmospheric Administration. <http://

oceanexplorer.noaa.gov/explorations/

05stepstones/logs/summary/summary.

html> 

Bullard SG, Lambert G, Carman MR, Byrnes J, 

Whitlatch RB, Ruiz G, Miller RJ, Harris 

L, Valentine PC, Collie JS, Pederson J, 

McNaught DC, Cohen AN, Asch RG, 

Dijkstra J, Heinonen K (2007) The 

colonial ascidian Didemnum sp. A: 

current distribution, basic biology and 

potential threat to marine communities of 

the northeast and west coasts of North 

America. Journal of Experimental Marine 

Biology and Ecology 342:99–108

Cairns SD (1981) Marine flora and fauna of the 

northeastern United States. Scleractinia. 

NOAA Tech. Rep NMFS Circ. 438:14 pp

Cairns SD (1992) Worldwide distribution of the 

Stylasteridae. Scientia Marina 56:125-

130

Cairns SD, Bayer FM (2005) A review of the genus 

Primnoa (Octocorallia: Gorgonacea: 

Primnoidae), with the description of two 

new species. Bulletin of Marine Science 

77:225-256
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

NORTHEAST

219

Cairns SD , Chapman RE (2001) Biogeographic 

affinities of the North Atlantic deepwater 

Scleractinia. Pages 30-57 In: Willison 

JHM, Hall J, Gass SE, Kenchington ELR, 

Butler M, Doherty P (eds.) Proceedings 

of the First International Symposium on 

Deep-Sea Corals. Ecology Action Center, 

Halifax, NS

Conkling PW, ed. (1995) From Cape Cod to the 

Bay of Fundy: an environmental atlas of 

the Gulf of Maine. Cambridge, MA: MIT 

Press. 272 pp

Cook SK (1988) Physical oceanography of the 

Middle Atlantic Bight. In: Pacheco, AL (ed.) 

Characterization of the Middle Atlantic 

water management unit of the Northeast 

regional action plan. NOAA Tech. Memo. 

NMFS-F/NEC-56:1-50

Deichman E (1936) The Alcyonaria of the western 

part of the Atlantic Ocean. Harvard Univ., 

Mus. Comparative Zool. Mem. 53:1-317

Dorsey EM (1998) Geological overview of the 

sea floor of New England. In: Dorsey, EM, 

Pederson, J (eds.) Effects of fishing gear 

on the sea floor of New England. MIT Sea 

Grant Publication 98-4:8-14

Eckelbarger K, Simpson A (2005) Taxonomic 

and reproductive studies of octocorals. 

Ocean Explorer/Explorations. National 

Oceanic and Atmospheric Administration/

Mountains in the Sea 2004/Mission 

Summary. <http://oceanexplorer.noaa.

gov/explorations/04mountains/logs/

summary/summary.html> 

Fosså JH, Mortensen PB, Furevik DM (2002) 

The deep-water coral Lophelia pertusa

in Norwegian waters: Distribution and 

fishery impacts. Hydrobiologia 471:1-12

Freiwald A (2002) Reef-forming cold-water 

corals. Pages 365-385 in Wefer G, Billet 

D, Hebbeln D, Jorgensen BB, Schluter 

M, Van Weering T (eds.) Ocean margin 

systems. Berlin, Germany: SpringerVerlag

Gass SE, Martin Willison JH (2005) An 

assessment of the distribution of deep-sea 

corals in Atlantic Canada by using both 

scientific and local forms of knowledge. 

Pages 223-245 in Freiwald A, Roberts JM 

(eds.), Cold-water corals and ecosystems. 

Springer-Verlag Berlin, Heidelberg

Hall-Spencer J, Allain V, Fosså JH (2001) 

Trawling damage to Northeast Atlantic 

ancient coral reefs. Proceedings of 

the Royal Society. Series B. Biological 

Sciences 269:507-511

Hecker B (1980) Scleractinians encountered 

in this study. Appendix C. In: Canyon 

Assessment Study. U.S. Department of 

Interior Bureau of Land Management, 

Washington, DC, No. BLM-AA551-CT8-

49

Hecker B (1990) Variation in megafaunal 

assemblages on the continental margin 

south of New England. Deep-sea 

Research 1 37:37-57

Hecker B, Blechschmidt G (1980) Final historical 

coral report for the canyon assessment 

study in the Mid- and North Atlantic 

areas of the U.S. outer continental 

shelf: epifauna of the northeastern 

U.S. continental margin. Appendix A. 

In: Canyon Assessment Study. U.S. 

Department of Interior Bureau of Land 

Management, Washington, DC, USA, No. 

BLM-AA551-CT8-49

Hecker B, Blechschmidt G, Gibson P (1980) 

Final report for the canyon assessment 

study in the Mid- and North Atlantic 

areas of the U.S. outer continental 

shelf: epifaunal zonation and community 

structure in three Mid- and North Atlantic 

canyons. In: Canyon Assessment Study. 

U.S. Department of Interior Bureau of 

Land Management, Washington, DC, No. 

BLM-AA551-CT8-49. pp 1-139

Hecker B, Logan DT, Gandarillas FE, Gibson 

PR (1983) Megafaunal assemblages 

in Lydonia Canyon, Baltimore Canyon, 

and selected slope areas. Pages 1-140 

In: Canyon and slope processes study: 

Vol. III, biological processes. Final report 

for U.S. Department of Interior, Minerals 

Management Service. No. 14-12-001-

29178

Heikoop JM, Hickmott DD, Risk MJ, Shearer 

CK, Atudorei V (2002) Potential climate 

signals from the deep-sea gorgonian coral 

Primnoa resedaeformis. Hydrobiologia 

471:117-124
NORTHEAST

220

STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

Houghton RL, Heirtzler JR, Ballard RD, Taylor 

PT (1977) Submersible observations 

of the New England Seamounts. 

Naturwissenschaften 64:348-355

ICES (2003) Report of the ICES Advisory 

Committee on Ecosystems, 2003. ICES 

Cooperative Research Report. 262. 229 

pp

ITIS (2006) Integrated Taxonomic Information 

System. <http://www.itis.usda.gov> 

Koslow JA, Gowlett-Holmes K, Lowry JK, 

O’Hara T, Poore GCB, Williams A (2001) 

Seamount benthic macrofauna off 

southern Tasmania: community structure 

and impacts of trawling. Marine Ecology 

Progress Series. 213:111-125 

Krieger KJ (2001) Coral (Primnoa) impacted 

by fishing gear in the Gulf of Alaska. 

Pages 106-116 In: Willison, JHM, Hall, J, 

Gass, SE, Kenchington, ELR, Butler, M, 

Doherty, P (eds.) Proceedings of the First 

International Symposium on Deep-Sea 

Corals. Ecology Action Center, Halifax, 

NS. pp 106-116

Krieger KJ, Wing BL (2002) Megafauna 

associations with deepwater corals 

(Primnoa spp.) in the Gulf of Alaska. In: 

Watling, L, Risk, M (eds.) Biology of cold 

water corals. Hydrobiologia 471:83-90

Langton RW, Langton EW, Theroux RB, Uzmann 

JR (1990) Distribution, behavior and 

abundance of sea pens, Pennatula 

aculeata, in the Gulf of Maine. Marine 

Biology. 107:463-469

Lazier A, Smith JE, Risk MJ, Schwarcz HP (1999) 

The skeletal structure of Desmophyllum 

cristagalli: the use of deep-water corals in 

sclerochronology. Lethaia 32:119-130

Leverette TL, Metaxas A (2005) Predicting habitat 

for two species of deep-water coral on 

the Canadian Atlantic continental shelf 

and slope. Pages 467-479 in Freiwald 

A, Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin, 

Heidelberg

MacIsaac K, Bourbonnais C, Kenchington 

E, Gordon Jr. D, Gass S (2001) 

Observations on the occurrence and 

habitat preference of corals in Atlantic 

Canada. Pages 58-75 In: Willison JHM, 

Hall J, Gass SE, Kenchington ELR, 

Butler M , Doherty P (eds.) Proceedings 

of the First International Symposium on 

Deep-Sea Corals. Ecology Action Center, 

Halifax, NS.

Malahoff A, Embley RW, Fornari-Daniel JJ (1982) 

Geomorphology of Norfolk and Washington 

canyons and the surrounding continental 

slope and upper rise as observed from 

DSRV Alvin. Chichester, U.K: John Wiley 

& Sons. Pages 97-111. In: Scrutton RA 

and Talwani M (eds.) The ocean floor; 

Bruce Heezan commemorative volume 

McDonough JJ, Puglise KA (2003) Summary: 

Deep-sea corals workshop. International 

Planning and Collaboration Workshop for 

the Gulf of Mexico and the North Atlantic 

Ocean. Galway, Ireland, January 16-17, 

2003. NOAA Tech. Memo. NMFS-F/SPO60. 51 pp

Moore JA, Vecchione M, Collette BB, Gibbons 

R, Hartel KE, Galbraith JK, Turnipseed 

M, Southworth M, Watkins E (2003) 

Biodiversity of Bear Seamount, New 

England Seamount chain: results of 

exploratory trawling. Journal of Northwest 

Atlantic Fishery Science 31:363-372

Moore JA, Vecchione M, Collette BB, Gibbons 

R, Hartel KE (2004) Selected fauna of 

Bear Seamount (New England Seamount 

chain), and the presence of “natural 

invader” species. Archive of Fishery and 

Marine Research 51:241-250

Mortensen PB (2000) Lophelia pertusa

(Scleractinia) in Norwegian waters: 

distribution, growth, and associated 

fauna. Deptartment of Fisheries and 

Marine Biology, Univ. Bergen, Norway

Mortensen PB, Buhl-Mortensen L (2004) 

Distribution of deep-water gorgonian 

corals in relation to benthic habitat 

features in the Northeast Channel (Atlantic 

Canada) Marine Biology 144:1223-1238
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

NORTHEAST

221

Mortensen PB, Buhl-Mortensen L, Gordon Jr. 

DC, Fader GBJ, McKeown DL, Fenton 

DG (2005). Effects of fisheries on deepwater gorgonian corals in the Northeast 

Channel, Nova Scotia (Canada). In: 

Barnes PW, Thomas JP (eds.) Benthic 

habitats and the effects of fishing. 

American Fisheries Society Symposium 

41:369-382

NEFMC [New England Fishery Management 

Council] (2004) Amendment 2 to the 

Monkfish Fishery Management Plan. 

Prepared by New England Fishery 

Management Council, Mid-Atlantic 

Fishery Management Council, NMFS. 

<http://nefmc.org/monk/index.htm>

Monkfish/Plan Amendments/2. 524 p 

Opresko B (1980) Taxonomic description of 

some deep-sea octocorals of the Mid and 

North Atlantic. Appendix B. In: Canyon 

Assessment Study. U.S. Department of 

Interior Bureau of Land Management, 

Washington, DC, No. BLM-AA551-CT8-

49

Packer D (2003) Northeast region. In: NOAA 

Fisheries, Office of Science & Technology 

(eds.) Our Living Oceans – Habitat. 

Unpublished draft manuscript, 16 

September 2003. pp 49-61

Puglise K, Brock R (2003) NOAA and deepsea corals: background, issues, and 

recommendations. Unpublished paper, 

NOAA, Silver Spring, MD 8 pp

Risk MJ, Heikoop JM, Snow MG and Beukens 

R (2002) Lifespans and growth patterns 

of two deep-sea corals: Primnoa 

resedaeformis and Desmophyllum 

cristagalli. In: Watling, L, Risk, M (eds.) 

Biology of cold water corals. Hydrobiologia 

471:125-131

Rogers AD (1999) The biology of Lophelia 

pertusa (Linnaeus 1758) and other deepwater reef-forming corals and impacts 

from human activities. Internat. Rev. 

Hydrobiologia 84:315-406

Shank T, Waller R, Cho W, Buckman K, L’Heureux 

S, Scheirer D (2006) Evolutionary and 

population genetics of invertebrates. 

Ocean Explorer/Explorations/North 

Atlantic Stepping Stones/Log. National 

Oceanic and Atmospheric Administration. 

<h t t p : / / o c e a n e x p l o r e r. n o a a . g o v /

e x p l o r a t i o n s / 0 5 s t e p s t o n e s / l o g s /

summary/summary.html>

Schmitz WJ, Wright WR, Hogg NG (1987) 

Physical oceanography. Pages 27-56. 

In: Milliman, JD, Wright, WR (eds) The 

marine environment of the U.S. Atlantic 

continental slope and rise. Boston, MA: 

Jones and Bartlett Publishers

Sherman K, Jaworski NA, Smayda TJ, eds. 

(1996) The northeast shelf ecosystem 

-- assessment, sustainability, and 

management. Cambridge, MA: Blackwell 

Science. 564 pp

Stevenson DK, Chiarella L, Stephan D, Reid R, 

Wilhelm K, McCarthy J, Pentony M (2004) 

Characterization of the fishing practices 

and the marine benthic ecosystems of the 

northeast U.S. shelf, and an evaluation of 

the potential effects of fishing on Essential 

Fish Habitat. NOAA Tech. Memo. NMFSNE-181

Stumpf RP, Biggs RB (1988) Surficial morphology 

and sediments of the continental shelf of 

the Middle Atlantic Bight. In: Pacheco 

AL (ed.) Characterization of the Middle 

Atlantic water management unit of the 

Northeast regional action plan. NOAA 

Tech. Memo. NMFS-F/NEC-56:51-72

Tendal OS (1992) The North Atlantic distribution 

of the octocoral Paragorgia arborea

(L, 1758) (Cnidaria, Anthozoa) Sarsia 

77:213-217

Theroux RB, Grosslein MD (1987) Benthic 

fauna. In: Backus, RH, Bourne, DW 

(eds.) Georges Bank. Cambridge, MA: 

MIT Press. pp 283-295

Theroux RB, Wigley RL (1998) Quantitative 

composition and distribution of the 

macrobenthic invertebrate fauna of 

the continental shelf ecosystems of 

the northeastern United States. NOAA 

Technical Report NMFS-140. 240 pp
NORTHEAST

222

STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

Townsend DW (1992) An overview of the 

oceanography and biological productivity 

of the Gulf of Maine. Pages: 5-26 In: 

Townsend, DW, Larsen, PF (eds) The 

Gulf of Maine. NOAA Coastal Ocean 

Prog. Reg. Synth. Ser. 1

Tucholke BE (1987) Submarine geology. In: 

Milliman, JD, Wright, WR (eds) The 

marine environment of the U.S. Atlantic 

continental slope and rise. Boston, MA: 

Jones and Bartlett Publishers. pp 56-113

Valentine PC, Uzmann JR, Cooper RA (1980) 

Geology and biology of Oceanographer 

submarine canyon. Marine Geology 38: 

283-312

Verrill AE (1862) Notice of a Primnoa from 

Georges Bank. Proc. Essex Inst., Salem, 

MA, USA, 3:127-129

Verrill AE (1878a) Notice of recent additions to 

the marine fauna of the eastern coast 

of North America. American Journal of 

Science and Arts Series 3, 16:207-215

Verrill AE (1878b) Notice of recent additions to 

the marine fauna of the eastern coast of 

North America, No. 2. American Journal 

of Science and Arts Series 3, 16:371-379

Verrill AE (1879) Notice of recent additions to 

the marine fauna of the eastern coast of 

North America, No. 5. American Journal 

of Science and Arts Series 3, 17:472-474

Verril AE (1884) Notice of the remarkable 

marine fauna occupying the outer banks 

of the southern coast of New England. 

American Journal of Science and Arts 

Series 3, 28:213-2

Verrill AE (1922) The Alcyonaria of the Canadian 

Arctic Expedition, 1913-1918, with a 

revision of some other Canadian genera 

and species. Reports from the Canadian 

Arctic Expedition 1913-18, vol VIII: 

molluscs, echinoderms, coelenterates, 

etc. Part G: Alcyonaria and Actinaria

Watling L (2005) Deep-sea octocorals as homes 

for other species. Ocean Explorer/

Explorations. National Oceanic and 

Atmospheric Administration/Mountains in 

the Sea 2004/Coral Commensals. <http://

oceanexplorer.noaa.gov/explorations/

04mountains/background/commensals/

commensals.html>

Watling L, Auster P (2005) Distribution of deepwater Alcyonacea off the northeast coast 

of the United States. Pages 259-264 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin, Heidelberg

Watling L, Auster P, Babb I, Skinder C, Hecker 

B (2003) A geographic database 

of deepwater alcyonaceans of the 

northeastern U.S. continental shelf and 

slope. Version 1.0 CD-ROM. National 

Undersea Research Center, University of 

Connecticut, Groton, USA

Watling L, Moore J, Auster P (2005) Mapping the 

distribution of octocorals and assessing 

the overall biodiversity of seamounts. 

Ocean Explorer/Explorations. National 

Oceanic and Atmospheric Administration/

Mountains in the Sea 2004/Mission 

Summary. <http://oceanexplorer.noaa.

gov/explorations/04mountains/logs/

summary/summary.html> 

Watling L, Simpson A, Mosher C (2006) 

Taxonomy and distribution of deep-sea 

octocorals. Ocean Explorer/Explorations/

North Atlantic Stepping Stones/Log. 

National Oceanic and Atmospheric 

Administration. <http://oceanexplorer.

noaa.gov/explorations/05stepstones/

logs/summary/summary.html> 

Watling L, Mosher C (2006) Commensalism in 

the deep sea: relationships of invertebrates 

to their octocoral hosts. Ocean Explorer/

Explorations/North Atlantic Stepping 

Stones/Log. National Oceanic and 

Atmospheric Administration. <http://

oceanexplorer.noaa.gov/explorations/

05stepstones/logs/summary/summary.

html> 

Wiebe PH, Backus EH, Backus RH, Caron 

DA, Gilbert PM., Grassle JF, Powers 

K, Waterbury JB (1987) Biological 

oceanography. Pages 140-201. In: 

Milliman JD, Wright WR (eds.) The 

marine environment of the U.S. Atlantic 

continental slope and rise. Boston, MA: 

Jones and Bartlett Publishers 

Wigley RL (1968) Benthic invertebrates of the 

New England fishing banks. Underwater 

Naturalist 5:8-13
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Higher Taxon Species Distribution Depth Range (m) Reference

Phylum CnidariaClass AnthozoaSubclass HexacoralliaOrder ScleractiniaFamily Caryophillidae Caryophyllia ambrosia 

ambrosia Alcock, 1898

Widespread (cosmopolitan) distribution; found on Bear Seamount

1487-2286 Cairns and Chapman 

2001; Moore et al. 2003

Caryophyllia ambrosia caribbeana Cairns, 1979

Endemic to western Atlantic 183-1646 Cairns and Chapman 2001

Dasmosmilia lymani(Pourtales, 1871)

Widespread (cosmopolitan) distribution. Continental slope south of New England, Lydonia Canyon, continental shelf between Baltimore and Hudson Canyons, in Baltimore Canyon, and between 100-200 m on the shelf south of Hudson Canyon and in the head of Hudson Canyon

37-366 Hecker 1980; Hecker et al. 

1983; Hecker 1990; Cairns and Chapman 2001; Guida (unpublished data)

Deltocyathus italicus(Michelotti, 1838)

Amphi-Atlantic with a disjunct distribution 403-2634 Cairns and Chapman 2001

Desmophyllum dianthus (Esper, 1794)

Widespread (cosmopolitan) distribution; found in several canyons (Corsair, Heezen, Lydonia, Oceanographer, Baltimore, Norfolk; near Hudson); continental slope on the southwestern edge of Georges Bank, between Veatch and Hydrographer Canyons; in the Mid-Atlantic on the slope between Lindenkohl Canyon on the south and Carteret Canyon on the north; in the Mid-Atlantic on the slope bounded by Toms Canyon to the south and Meys Canyon to the north; Bear Seamount

183-2250 Hecker 1980; Hecker 

and Blechschmidt 1980; Hecker et al. 1980, 1983; Malahoff et al. 1982; Cairns and Chapman 2001; Moore et al. 2003

Appendix 5.1. List of deep coral species found in the waters off the Northeastern United States. ** = distribution information based on studies or surveys of a particular area of the Northeast Region, not on overall distribution.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION Higher Taxon Species Distribution Depth Range (m) Reference Lophelia pertusa (L, 1758) Widespread (cosmopolitan) distribution; Oceanographer Canyon wall; Bear Seamount 146-1200; 700-1300 Hecker 1980; Hecker and Blechschmidt 1980; Hecker et al. 1980; Cairns and Chapman 2001; Moore et al. 2003 Solenosmilia variabilis Duncan, 1873 Widespread (cosmopolitan) distribution; Lydonia canyon; on the slope bounded by Toms Canyon to the south and Meys Canyon to the north 220-1383 Hecker 1980; Hecker et al. 1983; Cairns and Chapman 2001 Vaughanella margaritata (Jourdan, 1895) Endemic to northwestern Atlantic; Bear Seamount 1267 Cairns and Chapman 2001; Moore et al. 2003 Family Dendrophylliidae Enallopsammia profunda (Pourtales, 1867) Endemic to western Atlantic 403-1748 Cairns and Chapman 2001 Enallopsammia rostrata (Pourtales, 1878) Widespread (cosmopolitan) distribution 300-1646 Cairns and Chapman 2001 Family Flabellidae Flabellum alabastrum Moseley, 1873 Amphi-Atlantic with contiguous distribution; Canyons (Corsair, Heezen, Lydonia Oceanographer, Alvin, Baltimore, Norfolk) and slopes; Bear Seamount. some may be F. angularis or F. moseleyi 357-1977 Hecker 1980; Hecker and Blechschmidt 1980; Hecker et al. 1980, 1983; Cairns and Chapman 2001; Moore et al. 2003 Flabellum angulare Moseley, 1876 Amphi-Atlantic with contiguous distribution; see also F. alabastrum 2266-3186 Hecker 1980; Hecker and Blechschmidt 1980; Hecker et al. 1980, 1983; Cairns and Chapman 2001; Moore et al. 2003

Flabellum macandrewiGray, 1849

Amphi-Atlantic with contiguous distribution; see also F. alabastrum

180-667 Hecker 1980; Hecker 

and Blechschmidt 1980; Hecker et al. 1980, 1983; Cairns and Chapman 2001; Moore et al. 2003

Javania cailleti (Duch. & Mich., 1864)

Widespread (cosmopolitan) distribution; Lydonia, Oceanographer Canyons 

30-1809 Hecker 1980; Hecker 

et al. 1983; Cairns and Chapman 2001

Family Fungiacyathidae Fungiacyathus fragilis 

Sars, 1872

Widespread (cosmopolitan) distribution 412-460 Cairns and Chapman 2001
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Higher Taxon Species Distribution Depth Range (m) Reference

Family Rhizangiidae Astrangia poculata (Ellis & 

Solander, 1786)

Endemic to western Atlantic 0-263 Theroux and Wigley 1998; 

Cairns and Chapman 2001

Order AntipathariaFamily Antipathidae Leiopathes sp.** Near and on Bear Seamount 1643, 1754 Brugler 2005

Cirrhipathes sp.** Off Virginia 262 Smithsonian Institution 

collections

Subclass OctocoralliaOrder Alcyonacea Family Alcyoniidae Alcyonium digitatum

Linné, 1758

Watling and Auster 2005

Anthomastus agassiziiVerrill, 1922 **

Hard substrates from Corsair Canyon to Hudson Canyon; outcrops in Corsair Canyon; in Heezen, Lydonia, Oceanographer Canyons; on slope near Alvin Canyon; on slope on the southwestern edge of Georges Bank, between Veatch and Hydrographer Canyons; in MidAtlantic on slope flanked by Lindenkohl Canyon to south and Carteret Canyon to north and on slope bounded by Toms Canyon to south and Meys Canyon to north; Bear Seamount

750-1326 Hecker and Blechschmidt 

1980; Hecker et al. 1980, 1983; Opresko 1980; Valentine et al. 1980; Hecker 1990; Moore et al. 2003; Watling and Auster 2005

Anthomastus grandiflorusVerrill, 1878 **

Soft substrates, highest densities in canyons; found in Corsair, Heezen, Oceanographer Canyons; seen near Hudson Canyon, Toms Canyon, in Baltimore Canyon, in axis of Norfolk Canyon

700-2600 Hecker and Blechschmidt 

1980; Hecker et al. 1980; Opresko 1980; Watling and Auster 2005

Family Clavulariidae Clavularia modesta

(Verrill, 1874)

Watling and Auster 2005

Clavularia rudis (Verrill, 1922)**

Found in axis of Heezen, Lydonia, Oceanographer Canyons

750-1099 Hecker and Blechschmidt 

1980; Hecker et al. 1980; Opresko 1980; Watling and Auster 2005
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION Higher Taxon Species Distribution Depth Range (m) Reference Family Nephtheidae Capnella florida (Rathke, 1806)** Lydonia, Oceanographer, Baltimore Canyons; axis of Heezon Canyon; wall of Corsair Canyon; continental slope south of New England off Georges Bank 350-1500 Hecker and Blechschmidt 1980; Hecker et al. 1980; Opresko 1980; Hecker 1990; Watling and Auster 2005 Capnella glomerata (Verrill, 1869)** Several individuals found in Lydonia Canyon 200-561 Hecker et al. 1980; Opresko 1980; Watling and Auster 2005 Gersemia fruticosa** (Sars, 1860) Near and in deep portion of Hudson Canyon; at the mouth of Norfolk Canyon; seen near heads of Toms and Carteret Canyons (i.e., between Baltimore and Hudson Canyons) 600-3100 Hecker and Blechschmidt 1980; Opresko 1980; Watling and Auster 2005 Gersemia rubriformis (Ehrenberg, 1934) Watling and Auster 2005 Order Gorgonacea Family Acanthogorgiidae Acanthogorgia armata Verrill, 1878 ** Found in many canyons (Corsair, Lydonia, Oceanographer, Alvin, near Hudson, Norfolk, Baltimore); seen on boulders or outcrops 350-1300 Hecker and Blechschmidt 1980; Hecker et al. 1980; Opresko 1980; Malahoff et al. 1982; Watling and Auster 2005 Family Anthothelidae Anthothela grandiflora (Sars, 1856) ** Lydonia, Oceanographer, Baltimore Canyons 450-1150 Hecker et al. 1980; Opresko 1980; Watling and Auster 2005

Family Chrysogorgiidae Chrysogorgia agassizii

(Verrill, 1883)

Several individuals that may be C. agassiziifound in the vicinity of Hudson Canyon.

2150 Watling and Auster 2005

Iridogorgia pourtalesiiVerrill, 1883

Watling and Auster 2005

Radicipes gracilis (Verrill, 1884)

Watling and Auster 2005
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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Higher Taxon Species Distribution Depth Range (m) Reference

Family Isididae Acanella arbuscula

(Johnson, 1862) **

Found in Corsair, Heezen, Oceanographer Canyons; on slope near Alvin, Baltimore Canyons; in Mid-Atlantic on slope flanked by Lindenkohl Canyon to south and Carteret Canyon to north and on slope bounded by Toms Canyon to south and Meys Canyon to north; continental slope south of New England off Georges Bank; seen on soft substrates

600-2000 Hecker and Blechschmidt 

1980; Hecker et al.1980; Opresko 1980; Hecker 1990; Theroux and Wigley 1998; Watling and Auster 2005

Keratoisis grayi Wright, 1869

Watling and Auster 2005

Keratoisis ornata Verrill, 1878

Watling and Auster 2005

Lepidisis caryophylliaVerrill, 1883**

Bear Seamount? Moore et al. 2003; Watling 

and Auster 2005

Family Paragorgiidae Paragorgia arborea

(Linné, 1758) **

Found in Gulf of Maine, Georges Bank, and Canyons (Lydonia, Oceanographer, Baltimore, Norfolk); probably Bear Seamount

300-1100 Wigley 1968; Hecker 

and Blechschmidt 1980; Hecker et al. 1980; Opresko 1980; Theroux and Grosslein 1987; Theroux and Wigley 1998; Moore et al. 2003; Watling and Auster 2005 

Family Plexauridae Paramuricea grandis

Verrill, 1883 **

Found in Gulf of Maine and canyons from Corsair to near Hudson, seen in Corsair, Heezen, Oceanographer, Lydonia Canyons; on slope near Alvin Canyon; on slope on the southwestern edge of Georges Bank, between Veatch and Hydrographer Canyons; in MidAtlantic on slope flanked by Lindenkohl Canyon to south and Carteret Canyon to north and on slope bounded by Toms Canyon to south and Meys Canyon to north 

400-2200 Hecker and Blechschmidt 

1980; Hecker et al. 1980, 1983; Opresko 1980; Valentine et al. 1980; Watling and Auster 2005
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION Higher Taxon Species Distribution Depth Range (m) Reference Paramuricea placomus (Linné, 1758) ** Gulf of Maine Watling and Auster 2005 Paramuricea n. sp. Watling and Auster 2005 Swiftia casta (Verrill, 1883)** Bear Seamount? Moore et al. 2003; Watling and Auster 2005 Family Primnoidae Narella laxa Deichmann, 1936 Watling and Auster 2005 Primnoa resedaeformis Gunnerus, 1763) ** Found in Gulf of Maine, Georges Bank, and Canyons (Lydonia, Oceanographer, Baltimore, Norfolk); south to off Virginia Beach, VA; probably Bear Seamount 91-548 Hecker and Blechschmidt 1980; Hecker et al. 1980, 1983; Opresko 1980; Valentine et al. 1980; Theroux and Grosslein 1987; Theroux and Wigley 1998; Moore et al. 2003; Cairns and Bayer 2005; Watling and Auster 2005; Heikoop et al. 2002. Thouarella grasshoffi Cairns, 2006 Watling and Auster 2005 Order Pennatulacea Family Anthoptilidae Anthoptilum grandiflorum Newfoundland to Bahamas, Lousiana, Chile, Hawaii, Antarctica, N. Europe 274-3651 US NMNH collection, OBIS Anthoptilum murrayi Lydonia Canyon to Puerto Rico, Hawaii, Aleutians, Japan, W. Africa, N. Europe 430-2491 (1538 m min in NE US) US NMNH collection, OBIS

Family Kophobelemnidae Kophobelemnon 

stelliferum

Newfoundland to South Carolina, Japan, W. Africa, N. Europe

393-2199 (1330 m 

min in NE US)

US NMNH collection, OBIS

Kophobelemnon scabrum Nova Scotia to Virginia 1977-2249 US NMNH collectionKophobelemnon tenue Massachusetts to Virginia 2491-4332 US NMNH collection

Family Pennatulidae Pennatula aculeata Newfoundland to Virginia, California, Iberia, N. 

Africa

119-3316 US NMNH collection, 

OBIS

Pennatula grandis New Jersey, Bay of Biscay 1850-2140 US NMNH collection, 

OBIS

Pennatula borealis Newfoundland to North Carolina, California 219-2295 US NMNH collection, 

OBIS
STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION

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Higher Taxon Species Distribution Depth Range (m) Reference

Family Protoptilidae Distichoptilum gracile Nova Scotia to North Carolina, W. Africa, N. 

Eurpoe

1211-2844 (doubtful 

report at 59 m)

US NMNH collection, OBIS

Protoptilum abberans Nova Scotia to Virginia 1483-2359 US NMNH collectionProtoptilum carpenteri Massachusetts to North Carolina, W. Africa, N. 

Europe

1334-2194 US NMNH collection, 

OBIS

Family Scleroptilidae Scleroptilum gracile Massachusetts to Virginia 2513-4332 US NMNH collection

Scleroptilum grandiflorum Massachusetts to North Carolina, Panama, W. 

Africa

1502-2505 US NMNH collection, 

OBIS

Family Umbellulidae Umbellula guntheri Massachusetts to Virginia, Louisiana 2683-3740 (3166 m 

min in NE US)

US NMNH collection

Umbellula lindahlii Massachusetts to the Virgin Islands, Louisiana, 

Suriname, N. Europe, Indian O.

549-3338 (1538 m 

min in NE US)

US NMNH collection, OBIS

Family Virgulariidae Balticina finmarchica Newfoundland to Massachusetts, NC (doubtful), 

the Virgin Islands, Alaska

37-2249 (229 m min in 

NE US)

US NMNH collection

Stylatula elegans New York-Florida, Iberia 20-812 (51 m min in 

NE US)

US NMNH collection, OBIS

** Distribution information based on studies of a particular area in the Northeast Region, not an overall distribution.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE NORTHEASTERN US REGION SPECIES DISTRIBUTION OR LOCATION SCLERACTINIANS: Stony corals Dasmosmilia lymani Continental shelf between Baltimore and Hudson Canyons, in Baltimore Canyon, and between 100-200 m on the shelf south of Hudson Canyon and in the head of Hudson Canyon; soft substrates. Desmophyllum cristagalli Same as 

D. dianthus. Outcrops and underhangs at depths from 1000-1900 m. Seen on outcrops in Corsair Canyon. Found in 

Heezen Canyon. Seen in deeper parts of Lydonia Canyon, and on boulders or outcrops in Oceanographer Canyon, between 650-1600 m. Found on an outcrop near Hudson Canyon. Occasionally in axis of Norfolk Canyon.

Flabellum alabastrum

Canyons and slope from 600-2500 m; some may be F. angulare or F. moseleyi. Seen in Corsair Canyon. Found in Heezen and Oceanographer Canyons on soft substrate. Seen on deep continental slope near Alvin Canyon. Found on slope south of Baltimore Canyon. Found in deeper parts of the continental slope south of Norfolk Canyon and in axis of Norfolk Canyon on soft substrate. 

Lophelia prolifera

Same as L. pertusa. West wall of Oceanographer Canyon at 1100 m; dead rubble also found on wall at depths from 700-1300 m.

Solenosmilia variabilis Large colony recovered from the east flank of Lydonia Canyon.Javania cailleti One specimen recovered in axis of Oceanographer Canyon between 935-1220 m.ALCYONACEANS: Soft coralsAnthomastus grandiflorus

Soft substrates, highest densities in canyons. In the northern canyons found from 700-1500 m, southern canyons from 1500-2200 m; as deep as 2600 m. Found in Corsair, Heezen (west wall), and Oceanographer Canyons. Seen near Hudson Canyon, Toms Canyon, in Baltimore Canyon, and in axis of Norfolk Canyon. Frequently seen where a species of Pennatula was also common.

Anthomastus agassizii

Hard substrates from Corsair Canyon to Hudson Canyon from 750-1900 m. Seen on outcrops in Corsair Canyon. Found in Heezen Canyon. Seen in deeper parts of Lydonia Canyon. On boulders or outcrops in Oceanographer Canyon; 1057-1326 m. Seen on deep continental slope near Alvin Canyon. Seen near heads of Toms and Carteret Canyons (i.e., between Baltimore and Hudson Canyons).

Eunephthya fruticosa

Same as Gersemia fructicosa (?). Southern part of study area at depths from 2300-3100 m. Seen near Hudson Canyon around 2250-2500 m and at the mouth of Norfolk Canyon; populations found in deep portion of Hudson Canyon. Seen near heads of Toms and Carteret Canyons (i.e., between Baltimore and Hudson Canyons). Different form seen in Corsair and Heezen Canyons between 600-1200 m may be E. florida (see Opresko 1980) (Same as Capnella florida?)

Eunephthya florida

Same as Capnella florida (?). Found in Lydonia, Oceanographer, Baltimore Canyons, but only high abundances in Lydonia at 350-1500 m. Axis of Heezon Canyon between 1100-1200 m; wall of Corsair Canyon between 600-1000 m.

Eunephthya glomerata Same as Capnella glomerata (?). Several individuals found in Lydonia Canyon at 200 m and 562 m depth.Trachythela rudis

Same as Clavularia rudis. Axis of Heezen Canyon at 1100 m, Lydonia Canyon at 900 m, Oceanographer Canyon at 750 m and 900 m.

Appendix 5.2. Deep coral species discussed in Hecker and Blechschmidt (1980), Opresko (1980) (octocorals), and Hecker (1980) (scleractinians), as well as Hecker et al. (1980). Species names are listed exactly as stated in the literature.
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SPECIES DISTRIBUTION OR LOCATIONGORGONACEANS: GorgoniansParagorgia arborea

Lydonia Canyon 300-900 m, Oceanographer Canyon around 300-1100 m, axis of Baltimore Canyon 400 m and 500 m, Norfolk Canyon 400-600.

Anthothela grandiflora Found in Lydonia, Oceanographer, and Baltimore Canyons between 450-1149 m.Acanthogorgia armata

Found in many canyons from 600-2500 m depth. Seen on boulders or outcrops in Corsair and Oceanographer Canyons; found in Lydonia and Oceanographer Canyons between 400-1299 m. Seen on deep continental slope near Alvin Canyon. Found on an outcrop near Hudson Canyon. Found at 350 m in Baltimore Canyon. Occasionally in axis of Norfolk Canyon on exposed outcrops.

Paramuricea grandis

Found from Corsair Canyon to Hudson Canyon between 750-2150 m. Found on wall and axis of Oceanographer Canyon; found at depths between 400-1349 m in Lydonia and Oceanographer Canyons.

Paramuricea borealis

Same as 

P. grandis, perhaps also 

P. placomus (?). Found from Corsair Canyon to a site near Hudson Canyon at depths of 700-

2200 m on hard substrates. Seen on outcrops in Corsair Canyon. Found in Heezen Canyon. Seen in deeper parts of Lydonia Canyon. On boulders or outcrops in Oceanographer Canyon. Seen on deep continental slope near Alvin Canyon. Not seen in Norfolk Canyon.

Primnoa reseda Same as 

P. resedaeformis. Found in Lydonia Canyon at 560 m, in Baltimore Canyon at 450 m, and Norfolk Canyon at 400 m.

Acanella arbuscula

On soft substrates from 600-1300 m depth in the north and 1500-2000 m depth in the south. Seen in Corsair, Heezen, and Oceanographer Canyons. Found in Oceanographer Canyon between 1046-1191 m. Seen on deep continental slope near Alvin Canyon. On slope just south of Baltimore Canyon. Northern and southern forms may be different species.

Chrysogorgia agassizii Several individuals that may be 

C. agassizii were found at 2150 m in the vicinity of Hudson Canyon.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

STATE OF DEEP CORAL ECOSYSTEMS

IN THE U.S. SOUTHEAST REGION:

CAPE HATTERAS TO SOUTHEASTERN FLORIDA

Steve W. Ross1 and Martha S. Nizinski2

1

 UNC-Wilmington, Center for Marine Science

5600 Marvin Moss Ln.

Wilmington, NC 28409

*Currently assigned (through Intergovernmental 

Personnel Act) to:

US Geological Survey, Center for Coastal & 

Watershed Studies, St Petersburg, FL

2

 National Marine Fisheries Service National 

Systematics Laboratory

Smithsonian Inst., P.O. Box 37012, NHB, WC-57, 

MRC-153

Washington, DC 20013-7012

I. INTRODUCTION

Unique and productive deep coral habitats are 

found off the southeastern United States. This 

region may have the best developed, most 

extensive (Hain and Corcoran 2004) deep coral 

areas in U.S. waters. These deep reef systems 

have been largely ignored until recently, and this 

is partly due to their rugged bottom topography 

and the fact that they are usually overlain by 

extreme currents (i.e., Gulf Stream). Deep 

coral ecosystems face increasing threats world 

wide (Morgan et al. 2006; Roberts et al. 2006). 

Fisheries are expanding rapidly into deeper 

regions (Koslow et al. 2000; Roberts 2002), 

and hydrocarbon exploration and development 

are now also exploiting these depths. Two 

general deep coral habitats are reviewed for the 

southeastern U.S.: one is located along the shelf 

edge off east-central Florida, formed by the stony 

coral Oculina varicosa, and the second includes 

deeper water slope habitats dominated by the 

hard coral Lophelia pertusa (plus other corals 

and sponges) occurring off North Carolina and 

on the Blake Plateau off South Carolina through 

the Straits of Florida.

Deep coral habitats have been poorly studied, 

particularly in the western Atlantic. With the 

exception of the Oculina banks, references on 

deep corals off the southeastern U.S. are largely 

geological with a few biotic observations, mostly 

on invertebrates (Reed 2002a, 2002b; references 

in Sedberry 2001; Reed et al. 2006). Studies 

elsewhere revealed that deep reefs harbor 

extensive, species-rich invertebrate populations 

(Jensen and Frederiksen 1992; Rogers 1999; 

Buhl-Mortensen and Mortensen 2004). Fish 

studies related to the deep coral banks are rare. 

Our investigations of deep coral systems off 

the southeastern U.S. have revealed that many 

species of fishes (Ross and Quattrini 2007) 

and invertebrates are closely associated with 

this unique deep-reef habitat. Yet, it is unclear 

whether the deep coral habitat is essential to 

selected fishes or invertebrates or whether they 

occupy it opportunistically (see conflicting views 

in Auster 2005; Costello et al. 2005; Ross and 

Quattrini 2007). 

We review the deep coral ecosystems off the 

southeastern U.S. Deep coral research to 

date, coral distributions, associated faunal 

assemblages, threats to the corals, and 

management strategies in the U.S. Exclusive 

Economic Zone (EEZ) are briefly summarized. 

This chapter covers the region from Cape 

Hatteras, NC, to Key Biscayne, FL, and a depth 

range of 60 to about 5000 m (deeper depths 

vary with EEZ boundary). We emphasize corals 

inhabiting waters deeper than 200 m, which is the 

bathymetric range where most of the deep corals 

in this region occur; the Florida Oculina reefs (60-

100 m) are an exception.

History of Deep Coral Research off the 

Southeastern U.S.

Oculina Banks (<150 m)

Large reefs formed by Oculina varicosa are 

restricted in distribution to the shelf edge off 

east-central Florida. Although O. varicosa

forms substantial structures, the reefs were 

heavily fished, and are concentrated in shallower 

waters (60-120 m) than other deep corals (see 

below), these extensive reefs were not described 

until the 1960s (see reviews in Reed 2002a, 
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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2002b; Reed et al. 2005). The first publication 

mentioning these corals along the outer shelf off 

Florida reported results from seismic transects, 

dredging, and drop cameras (Macintyre and 

Milliman 1970). However, commercial fishermen 

and Florida scientists apparently knew of these 

reefs earlier (Reed et al. 2005). Macintyre and 

Milliman (1970) noted Oculina clumps and coral 

debris along the shelf break from northern Florida 

southward and especially from Cape Canaveral to 

Palm Beach where ridges in 70-90 m were usually 

capped with Oculina. Surveys of these reefs, 

using manned submersibles, began in the 1970s 

(Avent et al. 1977). Since then, reef monitoring, 

utilizing submersibles and ROVs, has continued 

intermittently (Reed et al. 2005). Additionally, 

side-scan sonar surveys were conducted, and 

multibeam mapping is currently ongoing on these 

coral banks (Reed et al. 2005; A.N. Shepard, 

NURC, unpublished data). A variety of studies 

have documented coral growth, distributions, 

and upwelling effects on coral growth (Reed 

1980, 1981, 1983), invertebrate communities 

associated with these reefs (Reed et al. 1982; 

Reed and Mikkelsen 1987), Oculina reproduction 

(Brooke and Young 2003, 2005), and fishes 

associated with the Oculina banks (Reed and 

Gilmore 1981; Gilmore and Jones 1992; Koenig 

et al. 2000, 2005; Reed et al. 2006).

These reefs were heavily fished during previous 

decades and incurred much damage and 

reduction in reef size due to impacts of fishing 

gear. Research continues, particularly in the 

zones protected by the South Atlantic Fishery 

Management Council (SAFMC). Funding is 

lacking, but habitat mapping, restoration, and 

monitoring are high priorities.

Deep-sea slope corals (>250 m)

Historically, deep coral research off the 

southeastern U.S. was temporally and spatially 

sporadic. Until recently deep coral research 

was often a by-product of non-coral projects. 

Figure 6.1. Southeastern United States regional report area, indicating general areas of 

Oculina varicosa reefs and the deeper coral (Lophelia mostly) habitats sampled by Ross 

et al. from 2000-2005 (red stars). The Stetson Bank (white box) is described in the text. 

Note that these areas do not represent all sites where deep (> 200 m) corals occur nor 

all sites visited by other researchers. See Reed et al. (2005, 2006) and Partyka et al. (in 

press) for additional deep coral sites in this region.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

The major studies that documented deep corals 

in the area are briefly reviewed (in roughly 

chronological order); this review is not intended 

to be inclusive.

The first report of deep corals from the Blake 

Plateau resulted from the 1880 collections of the 

steamer Blake (Agassiz 1888). These collections 

were poorly documented, and the bottom on the 

Blake Plateau was characterized as being hard 

and barren (Agassiz 1888). The research vessel 

Albatross collected corals on the Blake Plateau 

in 1886 using beam trawls and tangles. Some 

Lophelia specimens in those collections were 

deposited in the National Museum of Natural 

History (Smithsonian Institution), but were 

otherwise poorly documented. Squires (1959) 

noted several scleractinian species collected 

by dredge in 1954 off Palm Beach, FL in 686 

m. Cairns (1979) re-examined Squires’ coral 

collections and corrected identifications, resulting 

in the following species: Lophelia pertusa, 

Crispatotrochus (=Caryophyllia) squiresi, 

Enallopsammia profunda, and Tethocyathus 

variabilis.

An area of very rough topography containing 

deep corals was discovered on the Blake Plateau 

off South Carolina, resulting from surveys by 

depth sounder in the mid-1950’s (Stetson et 

al. 1962). However, confirmation that these 

features supported extensive coral habitat 

was not achieved until they were dredged and 

photographed in 1961 (Stetson 1961). Stetson 

et al. (1962) gave the first detailed accounting 

of this area now called the “Stetson Banks” 

(Figure 6.1), confirming the occurrence of two 

major species of hard corals, Lophelia pertusa

and Enallopsammia (=Dendrophyllia) profunda. 

They also reported species of Bathypsammia, 

Caryophyllia, and Balanophyllia as well as 

abundant alcyonarians. Additional details from 

the 1961 cruise, including locations of hundreds 

of coral mounds, were described by Stetson et 

al. (1969). 

Through the 1960s a series of geological papers 

based largely on precision echosounding data 

noted that numerous mounds, termed coral 

mounds, existed on the Blake Plateau and the 

Florida-Hatteras slope (e.g., Uchupi and Tagg 

1966; Uchupi 1967; Zarudzki and Uchupi 1968). 

Pratt (1968) presented one photograph of Lophelia

corals on the Blake Plateau (“Stetson Banks”). In 

1967, five manned submersible dives using the 

DSRV Alvin were made in an area west of the 

“Stetson Banks.” Two of these dives confirmed the 

occurrence of Enallopsammia (=Dendrophyllia) 

and Lophelia in the region (Milliman et al. 1967). 

Additionally, coral topped mounds (to 15 m high) 

were described from along the slope off Biscayne 

Bay, FL (around 700-825 m) (Neumann and Ball 

1970), based on 1967 Alvin dives.

Although corals were discovered on the Blake 

Plateau in the 1880s and investigated in the late 

1950s and early 1960s (Squires 1959; Stetson 

et al. 1962), such corals were not reported off 

North Carolina until the late 1960s. Based on 

seismic profiling, Uchupi (1967) first noted the 

occurrence of a coral mound off Cape Lookout, 

NC, which may be the same area illustrated 

(figure caption without comment) by Rowe and 

Menzies (1968). Rowe and Menzies (1969) later 

suggested that Lophelia sp. occurred off the 

Carolinas in “discontinuous banks” along the 450 

m contour, but gave no specific data. Menzies 

et al. (1973) vaguely referenced a “Lophohelia” 

bank off Cape Lookout, repeating a figure in Rowe 

and Menzies (1969), and presented a bottom 

photograph of a reef in 458 m. Cairns (1979) 

plotted a locality for Lophelia off Cape Lookout. 

Aside from Uchupi’s (1967) observations, the 

above North Carolina records mostly originated 

from a training cruise of the R/V Eastward (E-25-

66, I.E. Gray, chief scientist) during which a coral 

bank was photographed by drop camera (station 

E-4937, 475 m) and dredged (E-4933, 425 m) on 

30 June 1966. The Menzies et al. (1973, Figure 

4-4 B) photograph is from that cruise. This coral 

bank was discovered accidentally (independently 

of Uchupi 1967) as a result of constantly running 

the Eastward’s depth sounder (L.R. McCloskey 

and G.T. Rowe, pers. comm.). There were a 

few other short Eastward cruises to this area off 

Cape Lookout directed by Menzies, Rowe, Gray, 

or McCloskey, but no coral data were published. 

This Eastward station area was trawled and 

surveyed by sonar in May 1983 (R/V Delaware 

II cruise, S.W. Ross, chief scientist), but no hard 

bottom or coral were found. Coral mounds 

were located in this vicinity during an undersea 

survey using the Navy’s NR-1 nuclear research 

submersible (15-18 Nov 1993, K.J. Sulak and 

S.W. Ross, unpublished data). Three major coral 

areas have been located and studied off North 

Carolina (Ross and Quattrini 2007; Partyka et 

al. in press), and other mounds may exist. All 
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three areas off North Carolina were surveyed 

by multibeam sonar during October 2006 (Ross 

and Nizinski, unpublished data), revealing many 

mounds that had not been known. The slope off 

Cape Lookout appears to be the northern extent 

of deep corals in the southeast region.

Over the next three decades most studies around 

southeastern U.S. deep coral areas continued to 

be geological and generally not directed toward 

corals. Exceptions include Cairns (1979, 1981, 

2000, 2001a), who listed ranges for deep sea 

Scleractinia and azooxanthellate corals in this 

region, relying mostly on museum records. 

Neumann et al. (1977) described hard carbonate 

mounds in the eastern Straits of Florida off Little 

Bahama Bank that were covered in various 

corals (Lophelia and Enallopsammia) and other 

invertebrates. They coined the term “lithoherms” 

for these structures. In this same area in 1982, 

and also using Alvin, researchers collected and 

aged several “coral” species, indicating that 

these animals lived from several hundred up to 

1800 years (Griffin and Druffel 1989; Druffel et 

al. 1990, 1995). These corals have annual rings 

that contain a wealth of information about past 

climates, ocean productivity, and contamination. 

This significant discovery has vast implications for 

the scientific value of deep corals as proxies for 

climate change and recorders of environmental 

histories (Williams B et al. 2006; Williams et al. 

in press). Ayers and Pilkey (1981) documented 

several coral banks, collected corals, and dated 

some coral samples during a study of sediments 

of the Florida-Hatteras slope and inner Blake 

Plateau. Depending on location in a core, their 

dead coral samples ranged in age from 5,000 to 

44,000 years old. They dated a living specimen 

at 680 years old, but suggested that this age 

probably reflected the age of the carbon pool in 

the surrounding water. Pinet et al. (1981) also 

mapped coral banks overlapping the same area 

as Ayers and Pilkey (1981). Blake et al. (1987) 

briefly mentioned the presence of some soft 

and hard corals on the Blake Plateau. Many 

deep-reef locations were indicated by the U.S. 

Geological Survey sidescan sonar mapping 

(cruises in 1987) of the continental slope (EEZSCAN 87 Scientific Staff 1991); however, habitats 

were not verified in this large scale geological 

survey. Perhaps the first study to document the 

invertebrate community associated with deepcoral habitat in this region reviewed biozonation 

of lithoherms in the northeastern Straits of Florida 

(Messing et al. 1990). Genin et al. (1992) noted 

that sponges and gorgonians were common 

along the outer Blake escarpment (2624-4016 m) 

based on observations made during Alvin dives 

in 1980. They suggested that these communities 

were unusually dense for sites lacking sediment. 

Popenoe (1994) discussed the distribution and 

formation of coral mounds on the Blake Plateau 

and presented a few bottom photographs. Paull 

et al. (2000) surveyed deep-coral habitats off 

the Florida-Georgia border, dated parts of the 

structures, and suggested that such habitat was 

very common. Their dating indicated that some 

mounds may range from 18,000 to 33,000 years 

old. Popenoe and Manheim (2001) extensively 

reviewed geology, history, and habitats of the 

Blake Plateau around the area of the Charleston 

Bump, discussing various parameters that may 

control coral mound formation. Wenner and 

Barans (2001) described benthic habitats of the 

Charleston Bump area and noted some of the 

invertebrates and fishes occurring with deep 

corals. George (2002) discussed a coral habitat, 

dominated by Bathypsammia tintinnabulum, 

southeast of Cape Fear, NC (“Agassiz Coral 

Hills”) in 650-750 m. Apparently, the B. 

tintinnabulum used by Emilini et al. (1978) came 

from the collections noted by George (2002). 

A multibeam sonar survey of this site in 2006 

(Ross and Nizinski, unpublished data) revealed 

a flat bottom with no suggestion of coral mounds. 

Reed (2002a, 2002b; Reed et al. 2006) described 

several large areas of deep corals on the Blake 

Plateau and listed some of the fauna observed. 

As part of a SEAMAP bottom mapping project, 

data and reports to be examined for evidence 

of deep corals in this area were summarized by 

Arendt et al. (2003). This project was completed 

in 2006 and will be incorporated into the South 

Atlantic Fishery Management Council’s internet 

display.

Beginning in 2000 and continuing through the 

present, deep coral (or related habitat) research in 

the southeastern U.S. was stimulated by funding 

of studies through the NOAA Office of Ocean 

Exploration (see http://oceanexplorer.noaa.

gov/explorations) and supplemented by other 

sources. Teams lead by Principal Investigators 

S.D. Brooke, S.A. Pomponi, S.W. Ross, and G.R. 

Sedberry explored deep-coral banks throughout 

the southeast, mapping habitats, cataloging 

fauna, and conducting basic biological studies. 

A multi-investigator effort to create detailed 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

habitat classifications (Southeastern U.S. 

Deep-Sea Corals initiative, SEADESC) from 

past submersible dives in the area is underway 

(Partyka et al. in press). Future publications are 

forthcoming from the considerable data collected 

by these efforts.

II. GEOLOGICAL SETTING

Geology of the southeastern U.S. continental 

shelf, slope and rise has been well studied (see 

reviews in Pratt 1968; Avent et al. 1977; Schlee 

et al. 1979; Dillon and Popenoe 1988; Popenoe 

and Manheim 2001). The southeastern U.S. shelf 

and slope have been classified as a carbonate 

sedimentary province whose sediments are 

largely of terrigenous origin (Pratt 1968). The 

shelf edge (<200 m), including the region of the 

Florida Oculina banks, is marked by numerous 

topographic prominences of various origins 

(Macintyre and Milliman 1970; Avent et al. 

1977; Thompson and Gilliland 1980), and these 

provide substrate for attachment of a largely 

subtropical fauna (including corals). Unlike the 

Middle Atlantic Bight and Gulf of Mexico, major 

canyons that cut across the slope are missing 

in the southeast region. From central North 

Carolina northward the slope is particularly steep 

and characterized by a slump topography (Pratt 

1968). The Gulf Stream is and has been (since 

the early Tertiary) a dominant force shaping the 

bottom topology of the southeast region, and has 

scoured a steep channel along most of its length, 

often exposing hard substrates and creating a 

rugged topography.

The continental slope through much of the 

southeastern U.S. (central North Carolina to 

the Straits of Florida) is atypical of most U.S. 

slope configurations. It is unusually wide and 

is dominated by the Blake Plateau, a broad 

depositional feature formed by the Gulf Stream 

(400-1250 m depths, Popenoe and Manheim 

2001). The Blake Plateau exhibits two major 

topographic breaks, one on its western margin, the 

Florida-Hatteras Slope (shelf to about 600 m), and 

the Blake Escarpment (descending to about 4800 

m) on the southeastern margin (south of the Blake 

Spur). The eastern slope of the Blake Plateau 

north of the Blake Spur exhibits a more typical, 

less steep profile and grades into the Blake Ridge 

and Carolina Rise (Markl et al. 1970). The Blake 

Ridge contains extensive gas hydrate deposits 

and the only known methane seep community in 

this area (Borowski et al. 1997; van Dover et al. 

2003). The Blake Plateau is an extension of the 

Bahamian carbonate province, with carbonates 

being contributed by pteropod and Globigerina

material as well as corals and other invertebrates 

(Pratt and Heezen 1964). Numerous scarps, 

mounds, plateaus, and depressions occur in 

this region, and deep corals are common along 

the edges of the scarps and ridges and on the 

mounds (Stetson et al. 1962; Pratt and Heezen 

1964). One dominant topographic structure of 

the Blake Plateau is the Charleston Bump, which 

presents a partial barrier to Gulf Stream flow, 

deflects the Gulf Stream seaward, and is heavily 

scoured by this current, exposing hard substrates 

(Popenoe and Manheim 2001). Manganese 

pavements and nodules are abundant over the 

plateau (Brundage 1972). The western side of 

the Blake Plateau particularly has been heavily 

eroded (mostly during the Pleistocene) by the 

Gulf Stream, exposing numerous depressions, 

Cretaceous to Miocene aged hard substrata, and 

mounds (Uchupi 1967).

The southern part of this region (to off Key 

Biscayne, southern border for this report) is mostly 

within the Straits of Florida. This area is swept 

by the Florida current (part of the Gulf Stream 

system), and the bottom exhibits closely spaced 

valleys, ridges, mounds and lithoherms that are 

part of a carbonate platform (Malloy and Hurley 

1970; Neumann et al. 1977). The northwestern 

side of the Straits of Florida is bordered by the 

Miami Terrace, a carbonate platform in 200 to 

400 m that exhibits varied hardbottom topography 

(Reed et al. 2005). A variety of deep corals 

occur on hard substrates of the Straits of Florida 

(Neumann et al. 1977; Messing et al. 1990; Reed 

et al. 2005, 2006).

III. OCEANOGRAPHIC SETTING

The dominant oceanographic feature that shapes 

much of the geology and biology of the outer 

shelf and slope off the southeastern U.S. is the 

Gulf Stream current (or Florida Current). This 

well studied system influences bottom conditions 

even at continental slope depths by cutting off 

shelf sediment delivery to offshore areas (mostly 

between northern Florida to southern North 

Carolina), transporting sediments northward, 

and facilitating high current speeds (up to 
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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2.5 kn, 125 cm/s) on the bottom (Pratt 1968; 

Popenoe and Manheim 2001). The jet-like flow 

of the Gulf Stream has a surface width around 

80-150 km with a depth of 800-1200 m, fastest 

current speeds being near the surface center 

(Bane et al. 2001). The Gulf Stream provides 

a seasonally stable temperature (generally >27o

C) and salinity (generally >36 psu) regime for the 

outer shelf and slope. This current transports 

an increasingly massive volume of water as it 

moves northward (Bane et al. 2001). Inshore of 

the Gulf Stream (ca. <40 m depth) the physics of 

the continental shelf water column is dominated 

by tides, meteorology (e.g., winds, rainfall), and 

gravity waves (Pietrafesa 1983; Mathews and 

Pashuk 1986).

This complex current meanders (influenced in 

part by bottom topography) and produces eddies 

that spin away from the main current (Atkinson et 

al. 1985; Bane et al. 2001). As the Gulf Stream 

is deflected offshore away from the shelf edge, 

particularly off South Carolina (by the Charleston 

Bump) and off central North Carolina, nitrogenrich deeper water upwells onto the outer shelf, 

leading to localized areas of enhanced carbon 

production (Atkinson et al. 1982; Lee et al. 1991). 

Much of this carbon is subsequently transported 

offshore (Lee et al. 1991). As the Gulf Stream 

has varied in position since the Pleistocene, it 

may alternately uncover deep bottom substrata 

suitable for deep coral settlement or facilitate 

burial of coral mounds (Zarudzki and Uchupi 

1968).

Being on the shelf edge, the Florida Oculina

banks experience more temperate conditions. 

Bottom water temperatures can vary widely over 

short time scales (days to weeks) and on some 

banks can range from about 7° to 27° C (Avent et 

al. 1977; Reed 1981), being alternately washed 

by warm Gulf Stream or inshore waters and 

deeper, cold waters. The colder conditions are 

usually caused by upwelling, which also provides 

an increased amount of nutrients (Reed 1983). 

Bottom visibility on these banks is generally low, 

current speeds can be variable (sometimes >50 

cm sec-1), and sedimentation rates moderately 

high (15-78 mg cm-2 day-1) (Reed 2002b).

While the surface and upper water column 

oceanography beyond the shelf edge are fairly 

well studied, bottom conditions over most of the 

slope are not well known. Long term data are 

particularly lacking. High current speeds have 

been reported (see above), but currents can vary 

from near zero to >50 cm sec-1 over short time 

scales (pers. obs.). Bottom currents are more 

complex around coral mound or rocky features 

and are accelerated through valleys and over 

the tops of mounds/ridges (pers. obs.). Recent 

multibeam sonar mapping suggested that long 

term current scouring helped shape deep-coral 

mounds off North Carolina, but that conditions 

were different at the deeper Stetson area habitats 

(Ross and Nizinski, unpublished data). Bottom 

temperatures around southeastern U.S. deep 

coral habitats (370-780 m) ranged from 5.4° to 

12.3° C and salinities varied little from 35 psu 

(Ross and Quattrini 2007). Similar environmental 

data from southeastern U.S. deep coral habitats 

were reported by Reed et al. (2006). 

IV. STRUCTURE-FORMING DEEP 

CORALS OF THE SOUTHEASTERN

U.S.

The southeastern U.S. slope area, including 

the slope off the Florida Keys, has a unique 

assemblage of deep-water scleractinians 

(Cairns and Chapman 2001). The warm 

temperate assemblage identified by Cairns and 

Chapman (2001), encompassing nearly the 

same geographic range as that covered here, 

consists of about 62 species, four of which are 

endemic to the region. This group of corals was 

characterized by many free-living species, few 

species living deeper than 1000 m, and many 

species with amphi-Atlantic distributions. For the 

southeastern U.S., in areas deeper than 200 m, 

we report a similar assemblage, consisting of 57 

species of scleractinians (including 47 solitary 

and ten colonial structure-forming corals), four 

antipatharians, one zoanthid, 44 octocorals, one 

pennatulid, and seven stylasterids (Appendix 

6.1). Thus, the region contains at least 114 

species of deep corals (Classes Hydrozoa and 

Anthozoa). We note, however, that this list is 

conservative, and we expect that more species 

will be discovered in the region as exploration 

and sampling increase. Since solitary corals do 

not form reefs and are poorly known, we do not 

treat them in detail. Below we discuss the major 

structure-forming corals (Appendix 6.1) that most 

contribute to reef-like habitats in the southeastern 

U.S.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Figure 6.2. Selected photographs (May 2003) from 

the Florida Oculina HAPC. Photo credit: L. Horn, 

NOAA Undersea Research Center at UNCWilmington.

We hypothesize that high profile deep-coral reefs 

concentrate biota and enhance local productivity 

in ways similar to seamounts (Rogers 1994; 

Koslow 1997). The ridges and reef mounds, 

some rising over 100 m from open substrata, 

accelerate bottom currents which favors attached 

filter-feeding invertebrates and other biota. Thus, 

the growing reef alters the physics of the water 

column, enhancing the environment for continued 

coral growth and faunal recruitment (Genin et al. 

1986).

a. Stony Corals (Class Anthozoa, Order

 Scleractinia)

The dominant structure-forming coral on the 

southeastern U.S. outer shelf (<200 m) is 

Oculina varicosa (ivory tree coral). Although it 

occurs from Bermuda and North Carolina south 

through the Gulf of Mexico and the Caribbean in 

2-152 m depths, this coral only forms large reefs 

off east-central Florida, 27o 32’ N to 28o 59’ N, in 

70-100 m (Figure 6.1; Reed 2002b). The shallow 

water form of Oculina may have symbiotic 

zooxanthellae, but the deeper form does not. 
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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240

The deeper reefs are almost monotypic mounds 

and ridges which exhibit a vertical profile of 3-35 

m (Avent et al. 1977; Reed 2002b). Superficially, 

these structures (Figure 6.2) resemble the deep 

reefs formed by Lophelia pertusa. Despite cool 

temperatures, the shelf edge Oculina exhibit rapid 

growth, probably facilitated by regular upwellings 

of nutrient rich water (Reed 1983).

Lophelia pertusa, the major structure-forming 

coral in the deep sea, is the dominant scleractinian 

off the southeastern U.S. This species has 

a cosmopolitan distribution, occurring on the 

southeastern U.S. slope, in the Gulf of Mexico, 

off Nova Scotia, in the northeastern Atlantic, the 

South Atlantic, the Mediterranean, Indian Ocean 

and in parts of the Pacific Ocean over a depth 

range of 50 to 2170 m (Cairns 1979; Rogers 

1999). The 3380 m depth record off New York for 

L. pertusa reported by Squires (1959) was based 

on a misidentifed specimen (Cairns 1979). Coral 

habitats dominated by L. pertusa are common 

throughout the southeastern U.S. from about 370 

to at least 800 m depth.

Although Lophelia may occur in small scattered 

colonies attached to various hard substrata, it 

also forms complex, high profile features. For 

instance, off North Carolina, Lophelia forms what 

may be considered classic mounds that appear 

to be a sediment/coral rubble matrix topped with 

almost monotypic stands of L. pertusa (Figure 

6.3). Along the sides and around the bases of 

these banks are rubble zones of dead, gray coral 

pieces which may extend large distances away 

from the mounds. To the south sediment/coral 

mounds vary in size, and L. pertusa and other 

hard and soft corals populate the abundant hard 

substrata of the Blake Plateau in great numbers 

(Figures 6.4 and 6.5).

Data are lacking on how Lophelia coral banks in 

the southeastern U.S. are formed. Hypotheses 

for coral mound formation in the northeastern 

Atlantic were proposed (Hovland et al. 1998; 

Hovland and Risk 2003; Masson et al. 2003), 

but it is unclear how relevant these are off the 

southeastern U.S. The mounds off North Carolina 

and those in other locations off the southeastern 

U.S. (particularly east of south-central Florida) 

appear to be formed by successive coral growth, 

collapse, and sediment entrapment (Wilson 1979; 

Ayers and Pilkey 1981; Paull et al. 2000; Popenoe 

and Manheim 2001). Other coral formations in 

the area (especially on the Blake Plateau) seem 

to form by coral colonization of appropriate hard 

substrates, without mound formation by the 

corals. If bottom currents are too strong, mound 

formation may be prevented (Popenoe and 

Manheim 2001) because sediments cannot be 

trapped. Ayers and Pilkey (1981) suggested that 

Gulf Stream currents may erode coral mounds, 

and that present coral bank sizes may be related 

to historical displacements of that current. 

Assuming currents also carry appropriate foods, 

it may be that currents with variable speeds 

or at least currents of moderate speeds (fast 

enough to facilitate filter feeding but not too fast 

to prevent sediment entrapment) coupled with a 

supply of sediment are the conditions necessary 

to facilitate coral mound formation (Rogers 1999). 

Regardless of how coral formations are created, 

we agree with Masson et al. (2003) that elevated 

topography appears to be an important attribute 

for well developed coral communities.

Deep-coral reefs are fragile and susceptible to 

physical destruction (Fossa et al. 2002). It is 

estimated that these deep reefs may be hundreds 

to thousands of years old (Neumann et al. 1977; 

Wilson 1979; Ayers and Pilkey 1981; Mikkelsen 

et al. 1982; Mortensen and Rapp 1998); 

however, aging data are so limited (especially in 

the western Atlantic) that age of coral mounds in 

the western Atlantic is unclear. Recent drilling 

on coral mounds off Ireland indicated that these 

structures started forming over two million 

years ago and that formation was not related 

to hydrocarbon seeps (Williams T et al. 2006). 

While the genetic structure (gene flow, population 

relationships, taxonomic relationships) of Lophelia

in the northeastern Atlantic is being described 

(Le Goff-Vitry et al. 2004), such studies are just 

beginning in the western Atlantic (C. Morrison 

et al. unpublished data). Preliminary genetic 

results from the southeast region suggest that the 

population structure of L. pertusa is more diverse 

than expected (C. Morrison et al. unpublished 

data). Understanding the population genetics 

and gene flow will provide insights into coral 

biology, dispersal and distribution of deep corals 

off the southeastern U.S.

Although Lophelia is the dominant hard coral off 

North Carolina, other scleractinians contribute 

to the overall complexity of the habitat (Table 

6.1). Overall, species diversity of scleractinians 

increases south of Cape Fear, NC, but L. pertusa
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Figure 6.3. Depth sounder profiles and selected bottom views of the three North Carolina Lophelia coral 

banks. Top panel shows living (white) and dead (gray) corals near and on ridge tops. Middle panel also 

illustrates various live and dead corals as well as coral rubble (center photo). Bottom panel shows mixed corals, anemones, and conger eel (left photo), slope covered with anemones and corals (center photo), and the 

slender upright growth form of Lophelia (right photo). Habitat photographs do not correspond to a particular 

location on the depth sounder profiles. Profiles and photo credit: Ross et al. unpublished data.

Cape Lookout Lophelia Banks A

Cape Lookout Lopehlia Banks B

Cape Fear Lophelia Banks
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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242

Stetson area

Jacksonville Banks

South Canaveral

Figure 6.4. Depth sounder profiles and selected bottom views of coral (and sponge) habitats on the Blake 

Plateau south of North Carolina. Habitat photographs do not correspond to a particular location on the depth 

sounder profiles. Profiles and photo credit: Ross et al. unpublished data.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Figure 6.5. Examples of black (A,B; Leiopathes spp., 

possibly Leiopathes glaberrima), bamboo (D; Keratoisis spp., possibly K. ornata) and gold (E; Gerardia

sp.) corals commonly found on the Blake Plateau 

south of North Carolina. Other corals (C) have not 

yet been identified. Photo credit: Ross et al. unpublished data.

A B

C D

E

is still dominant. For example, the colonial corals 

Madrepora oculata and Enallopsammia profunda, 

rare off Cape Lookout, NC, are relatively common 

south of Cape Fear, NC. These hard corals tend 

not to occur singly or as species-specific mounds, 

but rather live on or adjacent to the Lophelia

mounds. A variety of solitary corals (Appendix 

6.1) are also found off the southeastern U.S. 

Individuals are often attached to coral rubble 

or underlying hard substrata. Most species 

appear to be either uncommon or rare. But, in 

some instances, particularly in the central portion 

of the region, local abundance can be high. 

For example, aggregations of Thecopsammia 

socialis and Bathypsammia fallosocialis carpet 

the bottom adjacent to reef habitat at study sites 

off South Carolina and northern Florida (Ross et 

al., unpublished data).

b. Black corals (Class Anthozoa, Order

 Antipatharia)

Black corals (Families Leiopathidae and 

Schizopathidae, ca. four species) are important 

structure-forming corals on the southeastern 

U.S. slope (Figure 6.5; Table 6.1). These corals 

occur locally in moderate abundances, but their 

distributions seem to be limited to the region 

south of Cape Fear, NC. Colonies may reach 

heights of 1-2 m. Black coral colonies, occurring 

singly or in small aggregations, may be observed 

either in association with hard coral colonies 

or as separate entities. Some of these living 
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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244

components of the deep reefs attain ages of 

hundreds to thousands of years (Williams B et 

al. 2006; Williams et al. in press; C. Holmes and 

S.W. Ross, unpublished data), and thus, along 

with gold corals, are among the oldest known 

animals on Earth. Black corals form annual 

or regular bands, and these bands contain 

important chemical records on past climates, 

ocean physics, ocean productivity, pollution, and 

data relevant to global geochemical cycles. An 

effort to investigate these geochemical data is 

underway by U.S. Geological Survey (C. Holmes 

and S.W. Ross).

c. Gold corals (Class Anthozoa, Order

 Zoanthidae)

Gerardia spp. colonies are found most often 

singly away from other coral structure, but these 

corals are also found associated with colonies of 

other structure-forming corals such as Lophelia 

pertusa, Keratoisis spp., or antipatharians 

(Leiopathes spp.). Very little is known about this 

group of organisms. They apparently exhibit slow 

growth, reaching ages of at least 1800 years old 

(Griffin and Druffel 1989; Druffel et al. 1995) and 

may be valuable in paleoecology studies. 

Table 6.1. Attributes of structure forming deep-sea corals of the southeastern United States. 

Taxa

ReefBuildingAbundance

Max 

Colony 

Size

Morphology

Associations 

with Other 

StructureForming 

Invertebrates

Colony 

Spatial 

Dispersion

Overall 

Structural 

Importance

Lophelia 

pertusa Yes High Large Branching Many Clumped High

Solenosmilia 

variabilis No Low Small Branching Many Clumped Low

Enallopsammia 

profunda No LowMedium

SmallMedium Branching Many Clumped LowMedium

Madrepora 

oculata No Low Small Branching Many Clumped Low

Oculina 

varicosa Yes High Large Branching Many Clumped High

Madracis 

myriaster No Low SmallMedium Branching Many Clumped Low

Leiopathes 

glaberrima No Medium Medium -Large Branching Many Solitary Medium

Bathypathes 

alternata No Low Medium -Large Branching Many Solitary Low

Keratoisis spp.

No Medium Medium 

-Large Branching Many Solitary Medium

Table Key

Attribute Measure

Reef-Building Yes/No

Relative Abundance Low/ Medium/ High

Size (width or height) Small (< 30cm)/ Medium (30cm-1m)/ Large (>1m)

Morphology Branching/ Non-branching

Associations None/ Few (1-2)/ Many (>2)

Spatial Dispersion Solitary/ Clumped

Overall Rating Low/ Medium/ High
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

d. Gorgonians (Class Anthozoa, Order

 Gorgonacea)

The gorgonians are by far the most diverse taxon 

on the southeastern U.S. slope represented 

by seven families, 17 genera, and 32 species 

(Appendix 6.1). The diversity of gorgonians 

increases dramatically south of Cape Fear, 

NC. Additional sampling is likely to increase 

the numbers of known species in this group for 

this region. To date, material we collected off 

Jacksonville, FL represented a newly described 

species (Thourella bipinnata Cairns 2006); 

the specimen of Chrysogorgia squamata also 

collected off Jacksonville represented the fifth 

known specimen of this species and increased 

our knowledge of its geographic range (previously 

known only from the Caribbean). 

Bamboo corals (Family Isididae, four species), 

possibly the best known members of this group 

because of their larger size and distinctive 

morphology, are also important structure-forming 

corals off the southeast region. (Figure 6.5; Table 

6.1). They occur locally in moderate abundances, 

and their distributions also seem to be limited to 

the region south of Cape Fear, NC. Colonies may 

reach heights of 1-2 m. Bamboo coral colonies 

occur either singly or in small aggregations and 

may be observed either in association with hard 

coral colonies or as separate entities.

e. True soft corals (Class Anthozoa, Order

 Alcyonacea)

Three families, Alcyoniidae, Nephtheidae, and 

Nidaliidae, comprise the Alcyonacea off the 

southeastern U.S. No family is speciose; total 

known diversity for this group is only six species 

(Appendix 6.1). The most abundant species 

observed in the region is Anthomastus agassizi, 

which is relatively abundant at sites off Florida. 

It is usually attached to dead Lophelia, but 

some individuals have also been observed on 

dermosponges and coral rubble. The majority 

of the alcyonacean species are smaller in size, 

both in vertical extent and diameter, than the 

gorgonians. Thus, these corals add to the overall 

structural complexity of the habitat by attaching 

to hard substrata such as dead scleractinian 

skeletons and coral rubble.

Stoloniferans, a suborder (Stolonifera) within 

the Alcyonacea, are represented by one family 

(Clavulariidae) off the southeast region. Six 

species from four genera have been reported 

from the region (Appendix 6.1). One species, 

Clavularia modesta, is widespread throughout 

the western Atlantic; the other five species are 

known from North Carolina southward to the 

Caribbean. 

f. Pennatulaceans (Class Anthozoa, Order

 Pennatulacea)

Little is known about pennatulids (sea pens) off 

the southeastern U.S. It is unlikely that this group 

contributes significantly to the overall complexity 

and diversity of the system. No sea pens have 

been observed during recent surveys (Ross et 

al., unpublished data) and based on museum 

records, only one species (Kophobelemnon 

sertum) is known in the region (Appendix 6.1).

g. Stylasterids (Class Hydrozoa, Order

 Anthoathecatae)

Although not found in great abundances, 

stylasterids (lace corals) commonly occur off the 

southeastern U.S. Seven species representing 

four genera have been reported from the region 

(Appendix 6.1). Individuals observed in situ are 

often attached to dead scleractinian corals or coral 

rubble. Abundance and diversity of stylasterids 

increase southward from the Carolinas. 

V. SPECIES ASSOCIATIONS WITH DEEP

CORAL COMMUNITIES

Oculina Banks (<150 m)

The fish community on the Florida shelf edge 

Oculina banks is typical of the southeastern U.S. 

shelf edge reef fauna (see review in Quattrini 

and Ross 2006). At least 73 species of fishes 

are known from the Oculina reefs (GOMFMC 

and SAFMC 1982; Koenig et al. 2005; Reed et 

al. 2006), and like the invertebrate community, 

this is a sub-tropically derived fauna. In recent 

years commercial fishing on these reefs has 

significantly depleted members of the snappergrouper complex and caused habitat destruction 

(Koenig et al. 2000, 2005). Some groupers, 

Mycteroperca microlepis (gag) and M. phenax

(scamp), use the reefs as spawning aggregation 

sites (Gilmore and Jones 1992); however, these 

have also been negatively impacted by habitat 

destruction (Koenig et al. 2000).

The Florida Oculina reefs support a diverse 

invertebrate fauna with mostly sub-tropical 

affinities (Figure 6.2). Densities of associated 
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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246

invertebrates rival those of shallow coral reef 

systems (see review in Reed 2002b). Avent et 

al. (1977) presented a preliminary list of benthic 

invertebrates dredged from some Oculina

mounds. Analysis of 42 small Oculina colonies 

yielded about 350 invertebrate species, including 

262 mollusc species (Reed and Mikkelson 1987), 

50 decapod crustacean species (Reed et al. 1982), 

47 amphipod species, 21 echinoderm species, 15 

pycnogoid species, and 23 families of polychaetes 

(Reed 2002b). The invertebrate community has 

been reduced by habitat destruction (Koenig et al. 

2000). Although Oculina habitats appear to have 

more associated mobile macroinvertebrates than 

deeper coral areas, large sponges and soft/horny 

corals are less abundant (Reed et al. 2006).

Deep-sea slope coral areas (>150 m, but most 

>300 m)

Deep coral habitat may be more important to 

western Atlantic slope species than previously 

known. Some commercially valuable deep-water 

species congregate around deep-coral habitat 

(Table 6.2). Various crabs, especially galatheoids, 

are abundant on the deep reefs, playing a role 

of both predator on and food for the fishes. 

Other invertebrates, particularly ophiuroids, 

populate the coral matrix in high numbers. On 

the relatively barren Blake Plateau, reefs (coral 

and hardgrounds) and surrounding coral rubble 

habitat seem to offer abundant shelter and food.

There are few deep-coral ecosystem references 

for the southeast region related to fishes, and 

those are generally qualitative (fishes neither 

collected nor counted) or fishes were not a specific 

target of the research (Popenoe and Manheim 

2001; Weaver and Sedberry 2001; Reed et al. 

2005, 2006). In the most detailed study of fishes 

to date, Ross and Quattrini (2007) identified 99 

benthic or benthopelagic fish species on and 

around southeastern U.S. deep-coral banks, 19% 

of which yielded new distributional data for the 

region. Additional publications resulting from their 

fish database documented the anglerfish 

fauna (Caruso et al. 2007), midwater fish 

interactions with the reefs (Gartner et al. in 

review), a new species of eel (McCosker 

and Ross in press), and a new species of 

hagfish (Fernholm and Quattrini in press). 

Although some variability in fish fauna 

was observed over this region, most of 

the deep-coral habitat was dominated by 

relatively few fish species (Table 6.2, Figure 

6.6). Many of these species are cryptic, 

being well hidden within the corals (e.g., 

Hoplostethus occidentalis, Netenchelys 

exoria, Conger oceanicus). Various reef 

habitats were characterized by Laemonema 

melanurum, L. barbatulum, Nezumia 

sclerorhynchus, Beryx decadactylus, 

and Helicolenus dactylopterus (Ross 

and Quattrini 2007). Nearby off reef 

areas were dominated by Fenestraja 

plutonia, Laemonema barbatulum, Myxine 

glutinosa, and Chlorophthalmus agassizi. 

Beryx decadactylus usually occurs in 

large aggregations moving over the reef, 

while most other major species occur as 

single individuals. The morid, Laemonema 

melanurum, is one of the larger fishes 

abundant at most sites with corals. This fish 

seems to rarely leave the prime reef area, 

while its congener L. barbatulum roams 

over a broader range of habitats. Although 

Helicolenus dactylopterus (Figure 6.6) can 

Scientific name

Common name 

 (if known) 

Myxinidae (mixed Myxine 

glutinosa and Eptatretus spp.) hagfishes

Scyliorhinus retifer chain dogfish

Scyliorhinus meadi

Cirrhigaleus asper roughskin dogfish

Dysommina rugosa

Synaphobranchus spp. cutthroat eels

Conger oceanicus* conger eel

Netenchelys exoria

Nezumia sclerorhynchus

Laemonema barbatulum shortbeard codling

Laemonema melanurum reef codling

Physiculus karrerae

Lophiodes beroe

Hoplostethus occidentalis western roughy

Beryx decadactylus* red bream

Helicolenus dactylopterus* blackbelly rosefish

Idiastion kyphos

Trachyscorpia cristulata Atlantic thornyhead

Polyprion americanus* wreckfish

Table 6.2. Dominant benthic fish species (in phylogenetic 

order) observed and/or collected during submersible dives 

(2000-2005) on or near southeastern U.S. Lophelia habitat 

based on Ross and Quattrini (2007). Asterisk (*) indicate 

commercially important species
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Figure 6.6 Photographs of some common fish species of the southeastern US deep (> 200 m) coral habitats 

Photographs credit: S.W. Ross.

Polyprion americanus (wreckfish) Beryx decadactylus (red bream)

Laemonema melanurum (reef codling)

Conger oceanicus (conger eel)

Helicolenus dactylopterus (blackbelly rosefish) Hoplostethus occidentalis (western roughy)

be common in all habitats, it occurs most often 

around structures. It is intimately associated with 

the coral substrate, and it is abundant around 

deep-reef habitat. Results (Ross and Quattrini 

2007) suggested that some of the fishes observed 

around the deep-coral habitats may be primary 

(obligate) reef fishes.

One of the most impressive biological aspects 

of these coral habitats (aside from the corals 

themselves) is the diverse and abundant 

invertebrate fauna (Table 6.3 and Reed et al. 

2006). Eumunida picta (galatheoid crab; squat 

lobster) and Novodinia antillensis (brisingid 

seastar) were particularly obvious (Figure 6.7), 

perched high on coral bushes to catch passing 

animals or filter food from the currents. One very 

different aspect of the North Carolina deep-coral 
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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248

potential for impacts to deep-sea ecosystems is 

of great concern because communities at these 

greater depths are not able to sustain heavy 

fishing pressures, as the general longevity of their 

species, slow growth, and low dispersal rates 

often prevent recovery from damaging impacts 

(Koslow et al. 2000; Roberts 2002; Cheung et 

al. 2007). A large portion of the Oculina banks 

was closed to fishing due to destruction of habitat 

and concern for conservation of corals and the 

associated fauna. There is concern that fisheries 

may soon target other deep-coral ecosystems in 

the region.

Fishing Effects

Major human induced damage to habitat and biota 

has been documented on the east-central Florida 

shelf edge, Oculina reef tract. Extensive damage 

to corals and fish stocks from fishing operations 

was reported (Coleman et al. 1999; Koenig et al. 

2000, 2005), including decreased numbers and 

biomass of corals, decreased amounts of coral 

habitat, and declining fish stocks. The primary fish 

targets (snapper, grouper, porgy) on the Oculina

reefs are also generally considered overfished 

throughout the waters off the southeastern U.S. 

(SAFMC unpublished data).

On the slope some commercially-exploited 

deep-water fishes, like Polyprion americanus

(wreckfish; Vaughan et al. 2001) and Helicolenus 

dactylopterus (blackbelly rosefish), utilize 

Lophelia habitat extensively (Ross and Quattrini 

2007). Swordfish have been observed along 

the deep reefs (Reed et al. 2006; Ross and 

Quattrini 2007). Other potentially exploitable 

species, such as royal red shrimps, rock crabs, 

golden crab, squid, bericiform fish species, 

and eels, are also associated with deep-coral 

habitats. Signs of past fishing effort (trash, lost 

gear) were observed on some banks, but the 

extent to which fishermen sample these areas is 

unknown; therefore, estimations of fishing impact 

(Table 6.4) are problematic. The potential for 

new deep-water fisheries on and around these 

banks is unknown. At this time our impression is 

that benthic fishing impacts to corals and benthic 

fishery species beyond 200 m in this region are 

minimal.

habitat compared to the rest of the southeast 

region is the massive numbers of the brittle star, 

Ophiacantha bidentata, covering dead coral 

colonies, coral rubble, and to a lesser extent, 

living Lophelia colonies (Figure 6.7). It is perhaps 

the most abundant macroinvertebrate on these 

banks and may constitute a major food source for 

fishes (Brooks et al. 2007). In places the bottom 

is covered with huge numbers of several species 

of anemones (Figure 6.7). The hydroid fauna is 

also rich with many species being newly reported 

to the area and some species being new to 

science (Henry et al. in press). The abundance of 

filter feeders suggests a food rich habitat. Various 

species of sponges, echinoderms, cnidarians 

(Messing et al. 1990) and crustaceans (Wenner 

and Barans 2001) also have been reported from 

deep-coral reefs off Florida, the northeastern 

Straits of Florida and the Charleston Bump region 

(Reed et al. 2006). Reed et al. (2006) provided a 

preliminary list of invertebrates, mostly sponges 

and corals, from some deep-coral habitats on the 

Blake Plateau and Straits of Florida; however, 

most taxa were not identified to species. Lack 

of data on the invertebrate fauna associated with 

deep corals is a major deficiency.

Although the invertebrate assemblage associated 

with northeastern Atlantic Lophelia reefs has 

been described as being as diverse as shallow 

water tropical coral reefs (e.g., Jensen and 

Frederickson 1992), data analysis of invertebrates 

associated with western Atlantic deep corals is too 

preliminary to speculate on the degree of species 

richness. Preliminary data on the invertebrate 

fauna (Nizinski et al. unpublished data) seem 

to indicate a faunal and habitat transition with 

latitude. In addition to changes in reef structure 

and morphology (see above), relative abundance 

within a single species decreases, overall species 

diversity increases, and numerical dominance 

between species decreases with decreasing 

latitude. In contrast to some fishes, the reef 

associated invertebrate assemblage appears to 

use deep reefs more opportunistically.

VI. STRESSORS ON DEEP CORAL 

ECOSYSTEMS OF THE 

SOUTHEASTERN U.S.

Very little direct information exists to evaluate 

the health or condition of deep-coral reefs along 

the coast of the southeastern U.S. However, the 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Table 6.3. Preliminary list of dominant benthic megainvertebrates observed or collected on or near 

southeastern U.S. deep coral habitats. Corals are listed separately in Appendix 6.1. References are 1= Nizinski 

et al. unpublished data, 2= Reed et al. 2006, 3 = Henry et al. in review. 

Effects Of Other Human Activities

Other anthropogenic activities could have 

negative impacts on deep-coral habitats, but 

presently most of these do not appear to be 

issues. Currently there is a federal moratorium on 

hydrocarbon exploration in this region, and there 

are no active offshore production operations. If 

this moratorium is lifted, the potential impacts to 

deep-coral habitat should be carefully considered. 

Cable laying could cause physical damage to 

coral habitat, but to date such damage has not 

been documented off the southeastern U.S. 

Construction of a proposed liquefied natural gas 

(LNG) terminal with associated benthic pipelines 

off south Florida could impact deep-coral habitat. 

Dominant Non-Coralline Invertebrate Taxa 

Phylum Porifera (Sponges)

Class Demospongiae 

multiple species1,2

Class Hexactinellida (glass sponges)

multiple species1,2 including 

Aphrocallistes beatrix1

Phylum Cnidaria

Class Hydrozoa (Hydroids)

multiple species (≥ 37 species)3

Class Anthozoa 

Order Actinaria (anemones)

multiple species including Actinaugi 

rugosa (Venus flytrap anemone)1

Order Zoanthidea (zoanthids)

multiple species1,2

Phylum Mollusca

Class Cephalopoda

Squids, Ilex sp.1

Octopus, multiple species1

Class Gastropoda

Coralliophila (?) sp.1

Phylum Annelida

Class Polychaeta (polychaetes)

multiple species including Eunice sp.1

Phylum Arthopoda

Subphylum Crustacea

Class Malacostraca

Order Decapoda

Infraorder Anomura

Family Chirostylidae (squat lobster)

Eumunida picta 1,2

Gastroptychus salvadori1

Uroptychus spp.1

Family Galatheidae (squat lobster) 

Munida spp.1

Munidopsis spp.1

Superfamily Paguroidea (hermit crabs 

and their relatives)

multiple species1

Infraorder Brachyura

Family Pisidae 

Rochinia crassa (inflated spiny 

crab)1

Family Geryonidae 

Chaceon fenneri (golden deepsea 

crab)1,2

Family Portunidae

Bathynectes longispina (bathyal 

swimming crab)1,2

Other taxa

Shrimps, multiple species1

Phylum Echinodermata

Class Crinoidea (crinoids)

multiple species1

Class Asteroidea (sea stars)

multiple species1,2

Order Brisingida (brisingid sea star)

Family Brisingidae

Novodinia antillensis1

Class Ophiuroidea (brittle stars)

multiple species1, including Ophiacantha 

bidentata1

Class Echinoidea (sea urchins)

Order Echinoida

Family Echinidae

Echinus gracilis1

E. tylodes1

Order Echinothurioida

Family Echinothuriidae

Hygrosoma spp.2

Order Cidaroida

Family Cidaridae

Cidaris rugosa1

Stylocidaris spp.2
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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Figure 6.7. Photographs of common invertebrates of the southeastern U.S. deep (>200 m) coral habitats. 

Photo credit: Ross et al. unpublished data.

wall of unidentified anemones Eumunida picta (squat lobster) on 

Lophelia coral

Novodinia antillensis (brisingid sea star)

Ophiacantha bidentata (brittle stars) intertwined 

within the Lophelia coral matrix and Echinus sp. 

top center.

Bathynectes longispinus 

(bathyal swimming crab)

Antedonidae (swimming crinoid)

Bottom disturbance through construction of 

offshore tanker ports may impact coral areas, 

especially off Florida where deep water is 

closer to shore. Construction of wind farms for 

energy production has been recently proposed 

for offshore areas. While these would likely 

be in waters shallower than those occupied by 

deep corals, designs for deeper water systems 

exist. Coral growth can keep up with a certain 

amount of sedimentation (Reed 2002b), but high 

rates of sedimentation are detrimental to corals 

(Rogers 1990). We are unaware of references 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

documenting sedimentation impacts to deep 

corals of the southeast region (except Oculina, 

Reed 2002b), and if they exist, most such impacts 

would usually not be anthropogenic. Active 

disposal activities (e.g., industrial, municipal, or 

military wastes) seem to be either rare or absent 

in deep waters of this region. There do not appear 

to be any deep coral harvesting activities off the 

southeastern U.S., although there is potential for 

this (GOMFMC and SAFMC 1982). Some mineral 

resources exist throughout the area (e.g., sand, 

manganese), but we are unaware of any current 

mining of these along the southeastern U.S. shelf 

edge or slope.

Climate change has not noticeably impacted 

southeastern U.S. deep corals. Impacts from 

rising ocean temperature to azooxanthellate 

deep corals would be different, but unknown, 

than those to shallow corals where zooxanthellae 

are expelled. Changes in sea level (increases) 

are likely to have little impact. However, climate 

changes that would impact the speed and direction 

of the Gulf Stream current or the overall North 

Atlantic conveyor system could have far reaching 

and difficult to predict impacts on deep corals. 

Changes in these currents could affect sediment 

transport, food delivery, dispersal mechanisms, 

as well as ambient temperature and salinity 

conditions. Ocean acidification from increased 

atmospheric CO2 is a recently identified potential 

impact to corals (Guinotte et al. 2006).

To date only one invasive species, the lionfish 

(Pterois volitans), has been documented from this 

area within a depth range to impact the Oculina

bank communities (Meister et al. 2005). While 

widespread and seemingly abundant, lionfish 

have not yet been reported from the Oculina

area. Their maximum reported depth off the 

southeastern U.S. is 99 m (Meister et al. 2005); 

thus, they are not expected at the deeper slope 

coral areas. 

VII. MANAGEMENT OF FISHERY 

RESOURCES AND HABITATS

All scleractinian and black corals off the 

southeastern U.S. are listed in Appendix II of the 

Convention on International Trade in Endangered 

Species of Wild Fauna and Flora (CITES). The 

purpose of this international agreement between 

governments is to ensure that international trade 

in specimens of wild animals and plants does not 

threaten their survival. Thus, CITES imposes 

restrictions on international trade in non-fossil 

corals.

The South Atlantic Fishery Management Council, 

in cooperation and collaboration with the National 

Marine Fisheries Service (NMFS), is responsible 

for management of habitat and most fishery 

resources in federal waters of the southeastern 

U.S. (see www.safmc.net). Management is 

executed through single species or species group 

fishery management plans. Plans that regulate 

the snapper/grouper complex, coastal pelagics, 

and dolphin/wahoo relate to species using the 

shelf edge Oculina banks. Fewer species are 

exploited in the deeper slope waters. Harvest 

of golden deep-sea crab (Chaceon fenneri) 

is regulated through a fishery management 

plan, and wreckfish are managed as part of the 

snapper/grouper complex. The SAFMC is moving 

from single species management toward an 

ecosystem-based approach which incorporates 

a broader appreciation of ecosystem interactions. 

Table 6.4. Potential fishing gear impacts to deep water corals in the southeastern United States. 

Gear Type Severity of 

Impact

Extent of 

Impact

Geographic Extent 

of Use in Region

Overall Rating of 

Gear Impact

Bottom Trawl High High Low High

Mid-water Trawl Low Low Low Low

Dredge High Medium Low Medium

Bottom-set Longline Medium Low Low Low

Bottom-set gillnet Medium Low Low Low

Traps or Pots Medium Low Low Low
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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252

Swordfish, tunas, sharks and billfishes are 

managed by the Highly Migratory Division of 

NMFS.

Although not applicable to deep corals in this 

region, other species (e.g., sea turtles, whales) 

are protected through such regulations as 

the Endangered Species Act and the Marine 

Mammal Protection Act. Sea turtles may occur 

on the Oculina banks; however, most of the slope 

deep-coral habitat is too deep for sea turtles and 

many marine mammals. In areas to be explored 

for hydrocarbons or mined for minerals within the 

EEZ, the Minerals Management Service (U.S. 

Dept. of Interior) requires geohazards surveys, 

including documentation of corals, and conducts 

environmental impact reviews of these activities.

Protection of coral habitat, including deep-water 

forms, in this region was established in a Coral, 

Coral Reef, and Live/Hardbottom Habitat Fishery 

Management Plan (FMP) under the MagnusonStevens Fishery Conservation and Management 

Act (GOMFMC and SAFMC 1982). This FMP 

summarized biological and other data on all 

corals off the southeastern U.S. and in the Gulf 

of Mexico. Additionally, optimum harvest of stony 

corals and sea fans throughout the waters off the 

southeastern U.S. was set at zero (collection for 

education and research purposes is permitted). 

The recent reauthorization of the MagnusonStevens Fishery Conservation and Management 

Reauthorization Act (P.L. 109-479) allows 

councils to designate zones for the protection 

of deep corals and requires research on and 

monitoring of deep coral habitats. The only deep 

coral protected area off the southeastern U.S., 

the Oculina Habitat Area of Particular Concern 

(HAPC), was described in GOMFMC and SAFMC 

(1982), but no other deep-coral areas were so 

designated. Designation of the Oculina banks as 

an HAPC became final in 1984, and use of bottom 

disturbing gear was prohibited (Reed 2002b). 

Over the next 14 years, these regulations were 

refined and expanded in a series of Amendments 

to the FMP. Increased protection of the Oculina

banks was granted in 1994, with a total fishing 

ban within the original HAPC. The HAPC was 

doubled in size in 2000, and the new expanded 

area is now closed to towed bottom gear. In 2004, 

the ban on fishing was extended indefinitely. 

No other deep-coral habitats are designated or 

fall within marine protected areas (MPAs), HAPCs 

or marine sanctuaries. No corals in the area are 

listed as Endangered or Threatened under the 

Endangered Species Act. If other deep-coral 

reefs prove to be important habitat with a unique 

fauna (as they seem to be), these reefs should be 

considered for protection as are the Oculina coral 

reefs. There are a variety of potential threats to 

the deep-coral habitats (see above). MPAs or 

HAPCs may be viable options for protecting 

these systems. However, considerable amounts 

and types of data, especially detailed maps, are 

critical for evaluating how and whether to protect 

deep-coral ecosystems (Miller 2001).

The SAFMC is currently evaluating management 

strategies for southeastern U.S. deep corals. 

Considering the needs of the SAFMC to evaluate 

and manage deep-water habitats in a timely 

manner, the brief, unpublished descriptions of 

southeastern U.S. deep-coral banks provided by 

Ross (2006) and Reed (2004) served as interim 

tools facilitating potential management options 

for deep-coral habitats. Based on these reports 

six large areas were recommended as deep coral 

HAPCs; these recommendations were modified 

in 2006 (Figure 6.8). These proposed HAPC 

areas are included in the current regional FMP 

and Ecosystem Plan (R. Pugliese, pers. comm.). 

A research plan is being prepared by a SAFMC 

committee to outline gaps in our knowledge and 

to address the immediate need for data pertaining 

to deep-coral habitats on the southeastern U.S. 

continental slope.

VIII. REGIONAL PRIORITIES TO 

UNDERSTAND AND CONSERVE DEEP

CORAL COMMUNITIES

Basic data are lacking for the majority of 

coral habitats >200 m. Recommendations 

below largely result from basic data needs. 

Considering their habitat value for deep-sea 

communities, their fragility, and a general lack 

of data, locating, describing, and mapping deep 

corals and conducting basic biological studies in 

these habitats are global and regional priorities 

(McDonough and Puglise 2003; Roberts and 

Hirshfield 2003; Puglise et al. 2005).

Recommendations

 Detailed mapping of the southeastern U.S. 

shelf edge and slope is critical to better 

understand these habitats and evaluate their 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

contributions to slope ecology. Such mapping 

is the foundation for most other research and 

management activities. Multibeam mapping 

should be conducted as soon as possible, 

especially in the depth range of 350-800 m. 

While this recommendation relates to the 

whole slope off the southeastern U.S., priority 

should be given to known coral sites and 

areas of suspected coral mounds.

 Of the many important ecological/biological 

studies that could be proposed, a broad 

trophodynamics study of coral banks and 

surrounding areas (whole water column) 

would provide the most impact for funds 

expended. Knowing the flow of energy in a 

system facilitates evaluation of anthropogenic 

impacts and allows predictions about the 

consequences of natural change. Such 

information is critical to ecosystem based 

management.

 Species composition and distributions of 

deep corals within the region require better 

documentation. Collection efforts for corals 

for identification by taxonomic experts, should 

be initiated. The overall deep-coral fauna is 

also poorly known. Better documentation of 

the whole living habitat matrix and associated 

fauna, as was done for Oculina reefs, is 

needed.

 Deep corals and the underlying mounds need 

to be aged. Accurate growth data on the 

major structure forming corals (e.g., Lophelia, 

Madrepora, bamboo and black corals) are 

Figure 6.8. Deep coral areas (red outlines) proposed for protection as Habitat Areas of Particular Concern by 

the South Atlantic Fishery Management Council.
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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254

critical to evaluate how banks are formed and 

their present status (accreting, eroding). This 

type of research may need to be coupled 

with local sedimentation and bottom current 

studies.

 Significant amounts of paleoclimate or 

paleoenvironmental data can be obtained 

from some coral species. Such studies 

should be pursued.

 Genetic studies should continue or be initiated 

for the major coral species and dominant 

associated fauna to examine taxonomic 

status, dispersal, relationships among coral 

banks, and community genetics.

 If protected areas are established for 

southeastern U.S. deep-coral banks, plans 

for long term monitoring, research, education, 

and enforcement should accompany this 

strategy. The SAFMC is developing such a 

plan. Funding should be made available to 

execute the plans.

 Any deep-water fisheries that currently exist 

or that develop on or near the deep-coral 

banks should be carefully monitored and 

regulated as deep-water fauna are highly 

vulnerable to over fishing, and the habitat is 

subject to permanent destruction.

IX. CONCLUSION

The southeast region contains a huge area of 

diverse deep-coral habitat. Rugged topography 

and hard substrata are common on the outer 

shelf edge and slope and this physical structure 

facilitates development of coral mounds and 

other coral habitats. However, detailed maps 

are lacking, and a major mapping effort must 

be initiated. Accurate maps are crucial to our 

understanding of the extent of this habitat, for 

planning research, and to our ability to manage 

deep-coral habitat. A recent multibeam mapping 

cruise (Ross and Nizinski unpublished data), 

covering most of the known North Carolina sites 

and portions of the Stetson banks, revealed 

numerous mounds (probably coral mounds), 

ridges, scarps, and depressions that were 

unknown. Based on these and other findings, 

it seems probable that the waters off the 

southeastern U.S. contains the greatest diversity 

and concentrations of reef building deep corals 

on the U.S. continental slope.

The three North Carolina Lophelia areas 

represent the northernmost deep-coral banks 

off the southeastern U.S. Significant deepcoral habitats are not apparent on the U.S. East 

coast again until north of Cape Cod. Because 

these banks seem to be a northern terminus for 

a significant zoogeographic region, they may 

be unique in biotic resources as well as habitat 

expression. The banks so far examined off North 

Carolina are different from much of the coral 

habitat to the south on the Blake Plateau. The 

North Carolina features are dominated by dense 

thickets of living L. pertusa that cover the tops 

and sides of the banks; the banks are surrounded 

by extensive coral rubble zones. Unlike areas to 

the south, the diversity of other corals is low. 

Southeastern U.S. deep-coral systems support 

a well developed community that appears to be 

faunistically different from surrounding non-reef 

habitats. The fish community on these deep reefs 

is composed of many species that do not (or at 

least rarely) occur off the reefs (Ross and Quattrini 

2007). Therefore, they may be considered 

primary reef fishes, in a way similar to those on 

shallow reefs. Many fish species thought to be 

rare and/or outside their reported ranges have 

been found on these reefs (Ross and Quattrini 

2007). Most likely these species only appeared 

to be rare because they occurred in areas that 

were difficult to sample by conventional means. 

Thus, these deep-coral habitats support a fish 

community that appears to be tightly coupled to 

the habitat and has essentially escaped detection 

until recently. Invertebrate communities are also 

very diverse and well developed; however, their 

associations with the reef habitat seem to be 

more opportunistic than is the case for certain fish 

species. However, invertebrate groups are poorly 

known on the slope reefs, and additional data are 

required from diverse habitats to evaluate habitat 

associations and allow comparisons with other 

ecosystems.

It is clear that the continental slope of the southeast 

region is important for corals and biodiversity. 

This is evidenced from the numerous new coral 

habitats discovered, the wide ranging extent and 

diversity of corals, numerous species from a 

variety of taxa newly recorded for the area, the 

many species new to science, and the fact that 
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

more fishes were recorded around these banks 

than any other deep-coral habitats worldwide. 

Some corals also provide important scientific data 

that will increase our understanding of climate 

and oceanographic changes. Some corals and/

or associated fauna (e.g., sponges) may have 

significant biomedical value. The overall impact 

of biodiversity in marine systems is significant 

(Worm et al. 2006), and while biodiversity of 

southeastern U.S. deep-coral systems is still 

poorly documented, these ecosystems are 

obviously a major component of regional slope 

ecology. Their protection, coupled with ongoing 

research, is necessary.

X. REFERENCES

Agassiz A (1888) Three cruises of the United 

States Coast and Geodetic Survey steamer 

“Blake” in the Gulf of Mexico, in the 

Caribbean Sea, and along the Atlantic coast 

of the United States, from 1877 to 1880. vol 

1. Bulletin of the Museum of Comparative 

Zoology at Harvard University14:1-314

Arendt MD, Barans CA, Sedberry GR, Van Dolah 

RF, Reed JK, Ross SW (2003) Summary 

of seafloor mapping and benthic sampling 

in 200-2000m from North Carolina through 

Florida. Final Rept Deep Water Mapping 

Project Phase II. SAFMC. Charleston, SC

Atkinson LP, Pietrafesa LJ, Hofmann EE (1982) 

An evaluation of nutrient sources to Onslow 

Bay, North Carolina. Journal of Marine 

Research 40:679-699

Atkinson LP, Menzel DW, Bush KA (eds) (1985) 

Oceanography of the Southeastern U.S. 

Continental Shelf. Coastal and Estuarine 

Sciences 2. American Geophysical Union, 

Washington, DC 156 pp

Auster PJ (2005) Are deep-water corals important 

habitats for fishes? Pages 747-760 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin Heidelberg

Avent RM, King ME, Gore RH (1977) 

Topographic and faunal studies of shelf-edge 

prominences off the central eastern Florida 

coast. Internationale Revue der Gesamten 

Hydrobiologie 62:185-208

Ayers MW, Pilkey OH (1981) Piston cores and 

surficial sediment investigations of the 

Florida-Hatteras slope and inner Blake 

Plateau. In: Popenoe P (ed) Environmental 

geologic studies on the southeastern Atlantic 

outer continental shelf. USGS Open File 

Rept 81-582-A, p 5-1-5-89
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

SOUTHEAST

256

Bane JM Jr, Attkinson LP, Brooks DA (2001) 

Gulf Stream physical oceanography at the 

Charleston Bump: deflection, bimodality, 

meanders, and upwelling. In: Sedberry GR 

(ed) Island in the Stream: oceanography 

and fisheries of the Charleston Bump. Amer 

Fish Soc Symposium 25, Bethesda, MD, p 

25-36

Bayer FM (2001) New species of Calyptrophora 

(Coelenterata: Octocorallia: Primnoidae) 

from the western part of the Atlantic Ocean. 

Proceedings of the Biological Society of 

Washington 114:367-380

Blake JA, Hecker B, Grassle JF, Brown B, Wade 

M, Boehm PD, Baptiste E, Hilbig B, Maciolek 

N, Petrecca R, Ruff RE, Starczak V, Watling 

L (1987) Study of Biological Processes 

on the U.S. South Atlantic Slope and 

Rise: Phase 2. US Department of Interior, 

Minerals Management Service, Atl OCS 

Region, Environmental Assessment Section. 

Herndon, VA, 414 pp + appendices

Borowski WS, Paull CK, Ussler W III (1997) Carbon 

cycling within the upper methanogenic zone 

of continental rise sediments: an example 

from the methane-rich sediments overlying 

the Blake Ridge gas hydrate deposits. 

Marine Chemistry 57:299-311

Brooke S, Young CM (2003) Reproductive 

ecology of a deep-water scleractinian coral, 

Oculina varicosa, from the southeast Florida 

shelf. Continental Shelf Research 23:847-

858

Brooke S, Young CM (2005) Embryogenesis and 

larval biology of the ahermatypic scleractinian 

Oculina varicosa. Marine Biology 146:665-

675

Brooks RA, Nizinski MS, Ross SW, Sulak KJ 

(2007) Frequency of sublethal injury in a 

deepwater ophiuroid, Ophiacantha bidentata, 

an important component of western Atlantic 

Lophelia reef communities. Marine Biology 

152:307-314

Brundage WL (1972) Patterns of manganese 

pavement distribution on the Blake Plateau. 

In: Horn DR (ed) Ferromanganese deposits 

on the ocean floor. National Science 

Foundation. Washington, DC, pp 221-250

Buhl-Mortensen L, Mortensen PB (2004) 

Symbiosis in deep-water corals. Symbiosis 

37:33-61

Cairns SD (1979) The deep-water scleractinia 

of the Caribbean Sea and adjacent waters. 

Studies on the fauna of Curacao and other 

Caribbean Islands 57(180), 341 pp

Cairns SD (1981) Marine flora and fauna of the 

Northeastern United States, Scleractinia. 

NOAA Technical Report NMFS Circular 438, 

14 pp

Cairns SD (1986) A revision of the Northwest 

Atlantic Stylasteridae (Coelenterata: 

Hydrozoa). Smithsonian Contributions to 

Zoology 418, 131 pp

Cairns SD (1997) A generic revision and 

phylogenetic analysis of the Turbinoliidae 

(Cnidaria: Scleractinia). Smithsonian 

Contributions to Zoology 591, 55 pp

Cairns SD (2000) A revision of the shallow-water 

azooxanthellate Scleractinia of the western 

Atlantic. Studies on the Natural History of 

the Caribbean Region 75, 240 pp

Cairns SD (2001a) A generic revision and 

phylogenetic analysis of the Dendrophylliidae 

(Cnidaria: Scleractinia). Smithsonian 

Contributions to Zoology 615, 75 pp

Cairns SD (2001b) Studies on western Atlantic 

Octocorallia (Coelenterata: Anthozoa). Part 

1: The genus Chrysogorgia Duchassaing 

& Michelotti, 1864. Proceedings of the 

Biological Society of Washington 114:746-

787

Cairns SD (2006) Studies on western Atlantic 

Octocorallia (Coelenterata: Anthozoa). 

Part 6: The genera Primnoella Gray, 1858; 

Thouarella Gray, 1870; Dasystenella

Versluys, 1906. Proceedings of the Biological 

Society of Washington 119:161-194

Cairns SD, Bayer FM (2002) Studies on 

western Atlantic Octocorallia (Coelenterata: 

Anthozoa). Part 2. The genus Callogorgia

Gray, 1858. Proceedings of the Biological 

Society of Washington 115:840-867 

Cairns SD, Bayer FM (2003) Studies on 

western Atlantic Octocorallia (Coelenterata: 
SOUTHEAST

257

STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Anthozoa). Part 3. The genus Narella Gray, 

1870. Proceedings of the Biological Society 

of Washington 116:617-648

Cairns SD, Bayer FM (2004a) Studies 

on western Atlantic Octocorallia 

(Coelenterata: Anthozoa). Part 4. The 

genus Paracalyptrophora Kinoshita, 1908. 

Proceedings of the Biological Society of 

Washington 117:114-139

Cairns SD, Bayer FM (2004b) Studies on 

western Atlantic Octocorallia (Coelenterata: 

Anthozoa). Part 5. The genera Plumarella

Gray, 1870; Acanthoprimnoa, n. gen.; and 

Candidella Bayer, 1954. Proceedings of the 

Biological Society of Washington 117:447-

487

 Cairns SD, Chapman RE (2001) Biogeographic 

affinities of the North Atlantic deep-water 

Scleractinia. In: Willison JHM, Hall J, Gass 

SE, Kenchington ELR, Butler M, Doherty P 

(eds) Proc. First Int Symposium on Deepsea Corals. Ecology Action Centre, Halifax, 

Nova Scotia, pp 30-57

Cairns SD, Hoeksema BW, van der Land J (1999) 

Appendix: List of extant stony corals. Atoll 

Research Bulletin 459:13-46

Caruso JH, Ross SW, Sulak KJ, Sedberry GR 

(2007) Deep-water chaunacid and lophiid 

anglerfishes (Pisces: Lophiiformes) off the 

south-eastern United States. Journal of 

Fisheries Biology 70:1015-1026

Cheung, WWL, Watson R, Morato T, Pitcher TJ, 

Pauly D (2007) Intrinsic vulnerability in the 

global fish catch. Marine Ecology Progress 

Series 333:1-12

Coleman FC, Koenig CC, Eklund A-M, Grimes 

CB (1999) Management and conservation of 

temperate reef fishes in the grouper-snapper 

complex of the southeastern United States. 

American Fisheries Society Symposium 

23:233-242

Costello MJ, McCrea M, Freiwald A, Lundalv T, 

Jonsson L, Brett BJ, van Weering TCE, de 

Haas H, Roberts JM, Allen D (2005) Role of 

cold-water Lophelia pertusa coral reefs as 

fish habitat in the NE Atlantic. Pages 771-

805 in Freiwald A, Roberts JM (eds.), Coldwater corals and ecosystems. SpringerVerlag Berlin Heidelberg

Dillon WP, Popenoe P (1988) The Blake Plateau 

Basin and Carolina Trough. In: Sheridan 

RE, Crow JA (eds) The Atlantic Continental 

Margin, U.S. The Geology of North America, 

vol 1-2. The Geological Society of North 

America, p 291-327

Druffel ERM, King LL, Belastock RA, Buesseler 

KO (1990) Growth rate of a deep-sea coral 

using 210Pb and other isotopes. Geochimica 

et Cosmochimica Acta 54:1493-1500

Druffel ERM, Griffin S, Witter A, Nelson E, 

Southon J, Kashgarian M, Vogel J (1995) 

Gerardia: bristlecone pine of the deep-sea? 

Geochimica et Cosmochimica Acta 59:5031-

5036

EEZ-SCAN 87 Scientific Staff (1991) Atlas of 

the U.S. exclusive economic zone, Atlantic 

continental margin. US Geological Survey 

Miscellaneous Investigations Series I-2054

Emiliani C, Hudson JH, George RY (1978) 

Oxygen and carbon isotope growth record 

in a reef coral from the Florida Keys and a 

deep-sea coral from Blake Plateau. Science 

202:627-629

Fernholm B, Quattrini AM (in press) A new 

species of hagfish (Myxinidae: Eptatretus) 

associated with deep-sea coral habitat in the 

western North Atlantic. Copeia

Fossa JH, Mortensen PB, Furevik DM (2002) 

The deep-water coral Lophelia pertusa in 

Norwegian waters: distribution and fishery 

impacts. Hydrobiologia 471:1-12

Gartner JV Jr, Sulak KJ, Ross SW, Necaise 

A-M (in review) Persistent near-bottom 

aggregations of mesopelagic animals along 

the North Carolina and Virginia continental 

slopes. Marine Biology
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

SOUTHEAST

258

Genin A, Dayton PK, Lonsdale PF, Spiess FN 

(1986) Corals on seamount peaks provide 

evidence of current acceleration over deepsea topography. Nature 322:59-61

Genin A, Paull CK, Dillon WP (1992) Anomalous 

abundances of deep-sea fauna on a rocky 

bottom exposed to strong currents. DeepSea Research 1 39:293-302

George RY (2002) Ben Franklin temperate 

reef and deep sea “Agassiz Coral Hills” 

in the Blake Plateau off North Carolina. 

Hydrobiologia 471:71-81

Gilmore RG, Jones RS (1992) Color variation 

and associated behavior in the epinepheline 

groupers, Mycteroperca microlepis (Goode 

and Bean) and M. phenax Jordan and Swain. 

Bulletin of Marine Science 51:83-103

Griffin S, Druffel ERM (1989) Sources of carbon 

to deep-sea corals. Radiocarbon 31:533-

543

Guinotte JM, Orr J, Cairns S, Freiwald A, Morgan 

L, George R (2006) Will human-induced 

changes in seawater chemistry alter the 

distribution of deep-sea scleractinian corals? 

Frontiers Ecology and Environment 4:141-

146

Gulf of Mexico Fishery Management Council and 

South Atlantic Fishery Management Council 

(1982) Fishery Management Plan Final 

Environmental Impact Statement for Coral 

and Coral Reefs. Charleston, SC

Hain S, Corcoran E (eds) (2004) 3. The status of 

the cold-water coral reefs of the world. Pages 

115-135. In: Wilkinson C (ed) Status of coral 

reefs of the world: 2004. vol 1. Australian 

Inst Mar Sci Perth, Western Australia

Henry L-A, Nizinski MS, Ross SW (in press) 

Diversity, distribution, and biogeography of 

hydroid assemblages collected from deepwater coral habitats off the southeastern 

United States. Deep-sea Research I

Hovland M, Mortensen PB, Brattegard T, Strass 

P, Rokoengen K (1998) Ahermatypic coral 

banks off mid-Norway: evidence for a link 

with seepage of light hydrocarbons. Palaios 

13:89-200

Hovland M, Risk M (2003) Do Norwegian deepwater coral reefs rely on seeping fluids? 

Marine Geology 198:83-96

Jensen A, Frederiksen R (1992) The fauna 

associated with the bank-forming deepwater 

coral Lophelia pertusa (Scleractinaria) on 

the Faroe Shelf. Sarsia 77:53-69

Koenig CC, Coleman FC, Grimes CB, Fitzhugh 

GR, Scanlon KM, Gledhill CT, Grace M 

(2000) Protection of fish spawning habitat 

for the conservation of warm-temperate reeffish fisheries of shelf-edge reefs of Florida. 

Bulletin of Marine Science 66:593-616

Koenig CC, Shepard AN, Reed JK, Coleman FC, 

Brooke SD, Brusher J, Scanlon KM (2005) 

Habitat and fish populations in the deepsea Oculina coral ecosystem of the Western 

Atlantic. In: Barnes PW, JP Thomas (eds) 

Benthic Habitats and the Effects of Fishing. 

American Fisheries Society Symposium 41, 

Bethesda, MD, p 795-805

Koslow JA (1997) Seamounts and the ecology 

of deep-sea fisheries. American Scientist 

85:168-176

Koslow JA, Boehlert GW, Gordon JDM, Haedrich 

RL, Lorance P, Parin N (2000) Continental 

slope and deep-sea fisheries: implications 

for a fragile ecosystem. ICES Journal of 

Marine Science 57:548-557

Le Goff-Vitry MC, Pybus OG, Rogers AD (2004) 

Genetic structure of the deep-sea coral 

Lophelia pertusa in the northeast Atlantic 

revealed by microsatellites and internal 

transcribed spacer sequences. Molecular 

Ecology 13:537-549

Lee TN, Yoder JA, Atkinson LP (1991) Gulf Stream 

frontal eddy influence on productivity of the 

southeastern U.S. continental shelf. Journal 

of Geophysical Research 96:22191-22205

Macintyre IG, Milliman JD (1970) Physiographic 

features on the outer shelf and upper slope, 

Atlantic continental margin, southeastern 

United States. Geological Society of America 

Bulletin 81:2577-2598
SOUTHEAST

259

STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Malloy RJ, Hurley R (1970) Geomorphology 

and geologic structure: Straits of Florida. 

Geological Society of America Bulletin 

81:1947-1972

Markl RG, Bryan GM, Ewing JI (1970) Structure 

of the Blake-Bahama outer ridge. Journal of 

Geophysical Research 75:4539-4555

Masson DG, Bett BJ, Billett DSM, Jacobs CL, 

Wheeler AJ, Wynn RB (2003) The origin 

of deep-water, coral topped mounds in the 

northern Rockall Trough, Northeast Atlantic. 

Marine Geology 194:159-180

Mathews TD, Pashuk O (1986) Summer and 

winter hydrography of the U.S. South Atlantic 

Bight (1973-1979). Journal of Coastal 

Research 2:311-336

McCosker JE, Ross SW (in press) A New 

Deepwater Species of the Snake Eel Genus 

Ophichthus (Anguilliformes: Ophichthidae) 

from North Carolina. Copeia

McDonough JJ, Puglise KA (2003) Summary: 

Deep-sea corals workshop. International 

planning and collaboration workshop for the 

Gulf of Mexico and the North Atlantic Ocean. 

Galway, Ireland, January 16-17, 2003. NOAA 

Tech Memo NMFS-SPO-60, 51 pp

Meister HS, Wyanski DM, Loefer JK, Ross 

SW, Quattrini AM, Sulak KJ (2005) Further 

evidence for the invasion and establishment 

of Pterois volitans (Teleostei: Scorpaenidae) 

along the Atlantic coast of the United States. 

SE Nat 4:193-206

Menzies RJ, George RY, Rowe GT (1973) 

Abyssal Environment and Ecology of the 

World Oceans. John Wiley and Sons, New 

York, USA

Messing CG, Neumann AC, Lang JC (1990) 

Biozonation of deep-water lithoherms and 

associated hardgrounds in the northeastern 

Straits of Florida. Palaios 5:15-33

Mikkelsen N, Erlenkeuser H, Killingley JS, Berger 

WH (1982) Norwegian corals: radiocarbon 

and stable isotopes in Lophelia pertusa. 

Boreas 11:163-171

Miller CA (2001) Marine protected area 

framework for deep-sea coral conservation. 

p 145-155. In: Willison JHM, Hall J, Gass 

SE, Kenchington ELR, Butler M, Doherty P 

(eds). Proc. First International Symposium 

on Deep-Sea Corals. Ecology Action Centre. 

Nova Scotia Museum. Halifax, Nova Scotia

Milliman JD, Manheim FT, Pratt RM, Zarudzki EFK 

(1967) Alvin dives on the continental margin 

off the southeastern United States July 2-13, 

1967. Woods Hole Oceanographic Institute 

Technical Report Ref No. 67-80

Mortensen PB, Rapp HT (1998) Oxygen and 

carbon isotope ratios related to growth line 

patterns in skeletons of Lophelia pertusa 

(L) (Anthozoa, Scleractinia): implications 

for determination of linear extension rates. 

Sarsia 83:433-446

Morgan LE, Tsao C-F, Guinotte JM (2006) Status 

of Deep Sea Corals in US Waters, with 

Recommendations for their Conservation 

and Management. Marine Conservation 

Biology Institute, Bellevue, WA, 63 pp

Neumann AC, Ball MM (1970) Submersible 

observations in the Straits of Florida: geology 

and bottom currents. Geological Society of 

America Bulletin 81:2861-2874

Neumann AC, Kofoed JW, Keller G (1977) 

Lithoherms in the Straits of Florida. Geology 

5:4–10

Partyka ML, Ross SW, Quattrini AM, Sedberry 

GR, Birdsong TW, Potter J, Gottfried S (in 

press) Southeastern United States Deep Sea 

Corals (SEADESC) Initiative: a collaborative 

effort to characterize areas of habitat-forming 

deep-sea corals. NOAA Tech Memo OER 1. 

Silver Spring, MD, 176 pp

Paull CK, Neumann AC, am Ende BA, Ussler 

W III, Rodriguez NM (2000) Lithoherms on 

the Florida-Hatteras slope. Marine Geology 

166:83-101

Pietrafesa LJ (1983) Shelfbreak circulation, 

fronts and physical oceanography: east and 

west coast perspectives. In: The shelfbreak: 

critical interface on continental margins. 

Society of Economic Paleontologists and 

Mineralogists Special Publication 33:233-

250
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

SOUTHEAST

260

Pinet PR, Popenoe P, Otter ML, McCarthy SM 

(1981) An assessment of potential geologic 

hazards of the northern and central Blake 

Plateau. In: Popenoe P (ed) Environmental 

geologic studies on the southeastern Atlantic 

outer continental shelf. USGS Open File 

Rept 81-582-A, pp 8-1-8-48

Popenoe P (1994) Bottom character map of the 

northern Blake Plateau. USGS Open File 

Rept 93-724

Popenoe P, Manheim FT (2001) Origin and 

history of the Charleston Bump-geological 

formations, currents, bottom conditions, and 

their relationship to wreckfish habitats on the 

Blake Plateau. In: Sedberry GR (ed) Island 

in the Stream: oceanography and fisheries 

of the Charleston Bump. American Fisheries 

Society Symposium 25, Bethesda, MD, p 

43-93

Pratt RM (1968) Atlantic continental shelf and 

slope of the United States - physiography 

and sediments of the deep-sea basin. U S 

Geological Survey Professional Paper529-

B, 44 pp

Pratt RM, Heezen BC (1964) Topography of 

the Blake Plateau. Deep-sea Research 1 

11:721-728

Puglise KA, Brock RJ, McDonough JJ III (2005) 

Identifying critical information needs and 

developing institutional partnerships to 

further the understanding of Atlantic deepsea coral ecosystems. Pages 1129-1140 in 

Freiwald A, Roberts JM (eds.), Cold-water 

corals and ecosystems. Springer-Verlag 

Berlin Heidelberg

Quattrini AM, Ross SW (2006) Fishes associated 

with North Carolina shelf-edge hardbottoms 

and initial assessment of a proposed marine 

protected area. Bulletin of Marine Science 

79:137-163

Reed JK (1980) Distribution and structure of 

deep-water Oculina varicosa coral reefs off 

central eastern Florida. Bulletin of Marine 

Science 30:667-677

Reed JK (1981) In situ growth rates of the 

scleractinian coral Oculina varicosa

occurring with zooxanthellae on 6-m reefs 

and without on 80-m banks. Proceedings of 

the 4th International Coral Reef Symposium 

2:201-206

Reed JK (1983) Nearshore and shelf-edge 

Oculina coral reefs: the effects of upwelling 

on coral growth and on the associated faunal 

communities. NOAA Symposium Series 

Undersea Research 1:119-124

Reed JK (2002a) Comparison of deep-water 

coral reefs and lithoherms off southeastern 

U.S.A. Hydrobiologia 471:57-69

Reed JK (2002b) Deep-water Oculina coral 

reefs of Florida: biology, impacts, and 

management. Hydrobiologia 471:43-55

Reed JK (2004) Deep-water coral reefs of Florida, 

Georgia and South Carolina: a summary 

of the distribution, habitat, and associated 

fauna. Unpublished Rept to South Atlantic 

Fishery Management Council, Charleston, 

SC, 71pp

Reed JK, Gilmore RG (1981) Inshore occurrence 

and nuptial behavior of the roughtail 

stingray, Dasyatis centroura (Dasyatidae), 

on the continental shelf, east central Florida. 

Northeast Gulf Science 5:1-4

Reed JK, Mikkelsen PM (1987) The molluscan 

community associated with the scleractinian 

coral Oculina varicosa. Bulletin of Marine 

Science 40:99-131

Reed JK, Gore RH, Scotto LE, Wilson KA (1982) 

Community composition, structure, aereal 

and trophic relationships of decapods 

associated with shallow- and deep-water 

Oculina varicosa coral reefs. Bulletin of 

Marine Science 32:761-786

Reed JK, Shepard AN, Koenig CC, Scanlon 

KM, Gilmore RG Jr (2005) Mapping, 

habitat characterization, and fish surveys 

of the deep-water Oculina coral reef Marine 

Protected Area: a review of historical and 

current research. Pages 443-465 in Freiwald 

A, Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg
SOUTHEAST

261

STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Reed JK, Weaver DC, Pomponi SA (2006) 

Habitat and fauna of deep-water Lophelia 

pertusa coral reefs off the southeastern U.S.: 

Blake Plateau, Straits of Florida, and Gulf of 

Mexico. Bulletin of Marine Science78:343-

375

Roberts JM, Wheeler AJ, Freiwald A (2006) Reefs 

of the Deep: the biology and geology of coldwater coral ecosystems. Science 312:543-

547

Roberts CM (2002) Deep impact: the rising toll of 

fishing in the deep sea. Trends In Ecology 

And Evolution 17:242-245

Roberts S, Hirshfield M (2003) Deep Sea Corals: 

out of sight, but no longer out of mind. 

Oceana, Washington, DC, USA , 16 pp

Rogers AD (1994) The biology of seamounts. 

Advances in Marine Biology 30:306-350 

Rogers AD (1999) The biology of Lophelia 

pertusa (Linnaeus 1758) and other deepwater reef-forming corals and impacts from 

human activities. International Review of 

Hydrobiology 84:315-406

Rogers CS (1990) Responses of coral reefs and 

reef organisms to sedimentation. Marine 

Ecology Progress Series 62:185-202

Ross SW (2006) Review of distribution, habitats, 

and associated fauna of deep water coral 

reefs on the southeastern United States 

continental slope (North Carolina to Cape 

Canaveral, FL). Unpublished Rept to South 

Atlantic Fishery Management Council, 

Charleston, SC. 2nd Ed, 37 pp

Ross SW, Quattrini AM (2007) The fish fauna 

associated with deep coral banks off the 

southeastern United States. Deep-Sea 

Research I 54:975-1007

Rowe GT, Menzies RJ (1968) Deep bottom 

currents off the coast of North Carolina. 

Deep-Sea Research 15:711-719

Rowe GT, Menzies RJ (1969) Zonation of large 

benthic invertebrates in the deep-sea off the 

Carolinas. Deep-Sea Research 16:531-537

Schlee JS, Dillon WP, Grow JA (1979) Structure 

of the continental slope off the eastern United 

States. In: Doyle, LJ, Pilkey OH (ed) Geology 

of Continental Slopes. Society of Economic 

Paleontologists and Mineralogists Special 

Publication 27, Tulsa, OK, pp 95-117

Sedberry GR (ed) (2001) Island in the Stream: 

oceanography and fisheries of the 

Charleston Bump. American Fisheries 

Society Symposium 25, Bethesda, MD, 240 

pp

Squires DF (1959) Deep sea corals collected 

by the Lamont Geological Observatory. I. 

Atlantic corals. American Museum Novitates 

No 1965:1-42

Stetson TR (1961) Report on Atlantis cruise # 266, 

June-July 1961. Woods Hole Oceanographic 

Institute Ref No 61-35

Stetson TR, Squires DF, Pratt RM (1962) Coral 

banks occurring in deep water on the Blake 

Plateau. American Museum Novitates 

2114:1-39

Stetson TR, Uchupi E, Milliman JD (1969) 

Surface and subsurface morphology of two 

small areas of the Blake Plateau. Trans Gulf 

Coast Assoc Geol Soc 19:131-142

Thompson MJ, Gilliland LE (1980) Topographic 

mapping of shelf edge prominences off 

southeastern Florida. Southeastern Geologyl 

21:155-164

Uchupi E (1967) The continental margin south 

of Cape Hatteras, North Carolina: shallow 

structure. Southeast Geology 8:155-177

Uchupi E, Tagg AR (1966) Microrelief of the 

continental margin south of Cape Lookout, 

North Carolina. Geological Society of 

America Bulletin 77:427-430

Van Dover CL, Aharon P, Bernhard JM, Caylor 

E, Doerries M, Flickinger W, Gilhooly W, 

Goffredi SK, Knick KE, Macko SA, Rapoport 

S, Raulfs EC, Ruppel C, Salerno JL, Seitz 

RD, Sen Gupta BK, Shank T, Turnipseed M, 

Vrijenhoek R (2003) Blake Ridge methane 

seeps: characterization of a soft-sediment, 

chemosynthetically based ecosystem. DeepSea Research I 50:281-300
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

SOUTHEAST

262

Vaughan DS, Manooch CS III, Potts JC (2001) 

Assessment of the wreckfish fishery on the 

Blake Plateau. pp 105-119 In: Sedberry GR 

(ed) Island in the Stream: oceanography and 

fisheries of the Charleston Bump. American 

Fisheries Society Symposium 25, Bethesda, 

MD

Weaver DC, Sedberry GR (2001) Trophic 

subsidies at the Charleston Bump: food web 

structure of reef fishes on the continental 

slope of the southeastern United States. 

pp 137-152 In: Sedberry GR (ed) Island in 

the Stream: oceanography and fisheries of 

the Charleston Bump. American Fisheries 

Society Symposium 25, Bethesda, MD

Wenner EL, Barans CA (2001) Benthic habitats 

and associated fauna of the upper- and 

middle-continental slope near the Charleston 

Bump. pp 161-178 In: Sedberry GR (ed) 

Island in the Stream: oceanography and 

fisheries of the Charleston Bump. American 

Fisheries Society Symposium 25, Bethesda, 

MD

Williams B, Risk MJ, Ross SW, Sulak KJ (2006) 

Deep-water Antipatharians: proxies of 

environmental change. Geology 34:773-776

Williams B, Risk MJ, Ross SW, Sulak KJ (in 

press) Stable isotope records from deepwater Antipatharians: 400-year records from 

the south-eastern coast of the United States 

of America. Bulletin of Marine Science

Williams T, Kano A, Ferdelman T, Henriet J-P, Abe 

K, Andres MS, Bjerager M, Browning EL, 

Cragg BA, De Mol B, Dorschel B, Foubert 

A, Frannk TD, Fuwa Y, Gaillot P, Gharib 

JJ, Gregg JM, Huvenne VAI, Leonide P, Li 

X, Mangelsdorf K, Tanaka A, Monteys X, 

Novosel I, Sakai S, Samarkin VA, Sasaki K, 

Spivack AJ, Takashima C, Titshack J. (2006) 

Cold-water coral mounds revealed. EOS 

87:525-526

Wilson JB (1979) “Patch” development of the 

deep-water coral Lophelia pertusa (L.) 

on Rockall Bank. Journal of the Marine 

Biological Association of the United Kingdom 

59:165-177

Worm B, Barbier EB, Beaumont N, Duffy JE, 

Folke C, Halpern BS, Jackson JBC, Lotze 

HK, Micheli F, Palumbi SR, Sala E, Selkoe 

KA, Stachowicz JJ, Watson R (2006) Impacts 

of biodiversity loss on ocean ecosystem 

services. Science 314:787-790

Zarudzki EFK, Uchupi E (1968) Organic reef 

alignments on the continental margin south 

of Cape Hatteras. Geological Society of 

America Bulletin 79:1867-1870
SOUTHEAST

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Appendix 6.1. Checklist of deep corals occurring off the southeastern United States (Cape Hatteras, NC to Key Biscayne, FL) at 200-1000 m depth (except shallower Oculina). Higher taxa are in phylogenetic order; families, genera and species are in alphabetical order. Some species have cosmopolitan distributions; however, only the northwestern Atlantic portion of their geographic ranges are reported. MR = museum records (holdings of the National Museum of Natural History, Smithsonian Institution). C & B = Cairns & Bayer (2002, 2003, 2004a, 2004b) *** = Cairns (1979, 2000), Cairns et al. 1999, Cairns & Chapman 2001, unpublished records. S = azooxanthellate solitary scleractinian corals. State of knowledge for the solitary corals is limited; therefore, speciesspecific geographic and bathymetric ranges are not given. Species included in this list have either been reported from the southeastern U.S. or are likely to occur in the region based on Cairns (1979, 2000), Cairns et al. 1999, and data obtained from unpublished records.

Higher Taxon Species Distribution Depth Range(m) ReferencePhylum CnidariaClass AnthozoaSubclass HexacoralliaOrder ScleractiniaFamily Anthemiphylliidae Anthemiphyllia patera Pourtalès, 1878

S

Family Caryophylliidae Anomocora fecunda (Pourtalès, 1871)

S

Asterosmilia marchadi (Chevalier, 1966)

S

Asterosmilia prolifera (Pourtalès, 1871)

S

Caryophyllia ambrosia caribbeana Cairns, 1979

S

Caryophyllia antillarum Pourtalès, 1874

S

Caryophyllia berteriana Duchassing, 1850

S

Caryophyllia polygona Pourtalès, 1878

S

Cladocora debilis Milne Edwards & Haime, 1849

S

Concentrotheca laevigata (Pourtalès, 1871)

S

Crispatotrochus squiresi (Cairns, 1979) 

S

Dasmosmilia lymani (Pourtalès, 1871)

S

Deltocyathus agassizii Pourtalès, 1867

S

Deltocyathus calcar Pourtalès, 1874

S

Deltocyathus eccentricus Cairns, 1979

S

Deltocyathus italicus (Michilotti, 1838)

S

Deltocyathus moseleyi Cairns, 1979

S

Deltocyathus pourtalesi Cairns, 1979

S
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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Higher Taxon Species Distribution Depth Range Reference

Desmophyllum dianthus (Esper, 1794)

S

Labyrinthocyathus facetus Cairns, 1979

S

Labyrinthocyathus langae Cairns, 1979

S

Lophelia pertusa (Linnaeus, 1758)

Nova Scotia - FL Straits; eastern Gulf of Mexico; Lesser Antilles

95-2000; 

commonly 500-

800 ***

Oxysmilia rotundifolia (Milne Edwards & Haime, 1848)

S

Paracyathus pulchellus (Philippi, 1842)

S

Premocyathus cornuformis (Pourtalès, 1868) 

S

Solenosmilia variabilis Duncan, 1873 GA - Suriname 220-1383 ***Stephanocyathus coronatus (Pourtalès, 1867)

S

Stephanocyathus diadema (Moseley, 1876)

S

Stephanocyathus laevifundus Cairns, 1977

S

Stephanocyathus paliferus Cairns, 1977

S

Tethocyathus cylindraceus (Pourtalès, 1868)

S

Tethocyathus recurvatus (Pourtalès, 1878)

S

Tethocyathus variabilis Cairns, 1979

S

Trochocyathus rawsonii Pourtalès, 1874

S

Family Dendrophylliidae Balanophyllia cyathoides (Pourtalès, 1871)

S

Balanophyllia floridana Pourtalès, 1868

S

Bathypsammia fallosocialis Squires, 1959

S

Bathypsammia tintinnabulum (Pourtalès, 1868)

S

Cladopsammia manuelensis (Chevalier, 1966)

Straits of FL; northern Gulf of Mexico; Arrowsmith Bank, Yucatan 55-366 ***

Eguchipsammia gaditana (Duncan, 1873)

NC, GA; Arrowsmith Bank, Yucatan 146-505 ***

Enallopsammia profunda (Pourtalès, 1867) MA - Straits of Florida 403-1748 ***Enallopsammia rostrata (Pourtalès, 1878) GA; off Nicaragua 300-1646 ***
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Higher Taxon Species Distribution Depth Range Reference

Thecopsammia socialis Pourtalès, 1868

S

Family Flabellidae Flabellum atlanticum Cairns, 1979

S

Flabellum moseleyi Pourtalès, 1880

S

Javania cailleti (Duchassaing & Michelotti, 1864)

S

Polymyces fragilis (Pourtalès, 1868)

S

Family Fungiacyathidae Fungiacyathus symmetricus (Pourtalès, 1871)

S

Family Guyniidae Pourtalocyathus hispidus (Pourtalès, 1878)

S

Schizocyathus fissilis Pourtalès, 1874

S

Stenocyathus vermiformis (Pourtalès, 1868)

S

Family Oculinidae Madrepora carolina (Pourtalès, 1871) 

NC - FL; Greater Antilles; western Caribbean; Gulf of Mexico

53-801; commonly 

200-300 ***

Madrepora oculata Linnaeus, 1758

GA - Rio de Janeiro, Brazil; Gulf of Mexico 144-1391 ***

Oculina varicosa Lesueur, 1821 NC - FL; West Indies; Bermuda 3-150 ***

Family Pocilloporidae

Madracis myriaster (Milne Edwards & Haime, 1849)

GA - Suriname; throughout the Caribbean and Gulf of Mexico 20-1220 ***

Family Turbinoliidae Cryptotrochus carolinensis Cairns, 1988

S

Deltocyathoides stimpsonii (Pourtalès, 1871)

S

Order AntipathariaFamily Leiopathidae Leiopathes glaberrima (Esper, 1788)

GA; FL; Gulf of Mexico (FL;AL; LA); Jamaica; Campeche Bank, Mexico; Venezuela 37; 220-685

20 MR; 

Ross et al. 

unpub.

Leiopathes spp.

Family Schizopathidae Bathypathes alternata Brook, 1889

SC; GA; FL; Yucatan Channel (off Arrowsmith Bank) 412-658

3 MR; 

Ross et al. 

unpub.

Parantipathes sp.

Order ZoanthidaeFamily Gerardiidae Gerardia spp.
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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Higher Taxon Species Distribution Depth Range Reference

Subclass OctocoralliaOrder AlcyonaceaFamily Alcyoniidae Anthomastus agassizi Verrill, 1922

Canada (Nova Scotia, Newfoundland); MA; DE; GA; FL; Bahamas 320-3186

28 MR; 

Ross et al. 

unpub.

Anthomastus grandiflorus Verrill, 1878

Canada (off Nova Scotia, Newfoundland); MA; VA; NC

137-457; 750-

2919 41 MR

Bellonella rubistella (Deichmann, 1936)

FL; Bahamas; Colombia; Venezuela; Trinidad; Tobago; Suriname; Dominican Republic; St. Lucia 24-329 25 MR

Family Clavulariidae Clavularia modesta (Verrill, 1874)

Canada (off Nova Scotia, Newfoundland); ME; MA; SC; GA; FL 29-861 63 MR

Scleranthelia rugosa (Pourtalès, 1867)

SC; Bahamas; Dominican Republic; Martinique 175-586 9 MR

Telesto fruticulosa Dana, 1846 NC; SC; GA; FL 13-105 218 MRTelesto nelleae Bayer, 1961

NC; Straits of FL (off Havana, Cuba); Bahamas

27-298; 1023-

1153 19 MR

Telesto sanguinea Deichmann, 1936

SC; FL; Gulf of Mexico (off FL, LA) 24-134 44 MR

Trachythela rudis Verrill, 1922 FL 805 1 MR

Family Nephtheidae Gersemia fruticosa (Sars, 1890)

Canada (off Nova Scotia, Newfoundland); MA; DE; VA; FL

91-368; 770; 

2107-3506 42 MR

Pseudodrifa nigra (Pourtalès, 1868)

SC; GA; FL; Bahamas; Straits of FL (off FL Keys; Havana, Cuba); Gulf of Mexico (off FL Keys)

60-878; 1153-

1023

47 MR; 

Ross et al. 

unpub.

Family Nidaliidae Siphonogorgia agassizii (Deichmann, 1936) FL; Gulf of Mexico (FL; TX) 14-159; 350-400 27 MROrder GorgonaceaFamily Chrysogorgiidae Chrysogorgia multiflora Deichmann, 1936

GA; FL; Bahamas; Straits of FL (off Key West); Gulf of Mexico (FL Keys); Lesser Antilles; Brazil 320-1354

Cairns, 

2001b; 14 

MR

Chrysogorgia squamata (Verrill,1883) Jacksonville, Fl; Caribbean 430-1050

Ross et al. 

unpub.
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Higher Taxon Species Distribution Depth Range Reference

Family Coralliidae Corallium sp.

Corallium niobe Bayer, 1964 off Jupiter Inlet; Bahamas 659-677; 1023 3 MR

Family Gorgoniidae Eunicella modesta (Verrill, 1883)

Savannah lithoherms, GA; east coast FL Lophelia reefs 518-732

Reed 2004; 

Ross et al. 

unpub.

Family Isididae Acanella eburnea (Pourtalès, 1868)

Hudson Canyon; SC; FL; Bahamas; Gulf of Mexico (FL; LA; TX); Caribbean Sea (Nevis) 309-2100 34 MR

Keratoisis flexibilis (Pourtalès, 1868)

GA; FL; eastern Gulf of Mexico; Bahamas; Campeche Bank, Mexico; Guadeloupe; Colombia; Venezuela 170-878 29 MR

Keratoisis ornata Verrill, 1878

Canada (off Nova Scotia, Newfoundland); MA; GA; FL; Bahamas; Cuba 274-3236

45 MR; 

Ross et al. 

unpub.

Lepidisis longiflora Verrill, 1883 FL; Caribbean Sea (Nevis) 743-1125 2 MR

Family Paragorgiidae Paragorgia arborea (Linnaeus, 1782) Canada; MA; NJ; MD; VA; NC 247-680 9 MR

Paragorgia johnsoni Gray, 1862

Florida Straits (off Palm Beach); Bahamas 522-608 6 MR

Family Plexauridae Paramuricea placomus (Linnaeus, 1758)

Canada (off Nova Scotia); GA; FL; Straits of FL (off Havana, Cuba) 247-805 6 MR

Paramuricea sp.Swiftia casta (Verrill, 1883)

MA; SC; GA; FL; Straits of FL (off FL Keys; Havana, Cuba); Bahamas; Gulf of Mexico (FL; LA); Yucatan Channel (off Arrowsmith Bank) 40-1953 49 MR

Swiftia exserta (Ellis & Solander, 1786)

GA; FL; Straits of FL (off FL Keys); Bahamas; Puerto Rico; Gulf of Mexico (FL; MS); Mexico; Panama; Colombia; Venezuela; Tobago; French Guiana; Guyana; Brazil 18-494 54 MR
STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

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Higher Taxon Species Distribution Depth Range Reference

Swiftia koreni (Wright & Studer, 1889)

Lydonia Canyon; FL; Gulf of Mexico (off FL Keys) 221-858 3 MR

Family Primnoidae

Callogorgia americana americana Cairns & Bayer, 2002 Straits of Florida; Lesser Antilles 183-732 C & B, 2002Callogorgia gracilis (Milne Edwards & Haime, 1857)

off central Florida; Bahamas; Antilles; off Honduras; northern Gulf of Mexico 82-514 C & B, 2002

Calyptrophora gerdae Bayer, 2001 Straits of FL 229-556 Bayer, 2001Calyptrophora trilepis (Pourtalès, 1868) SC; GA; Bahamas 593-911

Bayer, 

2001; 8 MR

Candidella imbricata (Johnson, 1862)

New England seamounts; Bermuda; eastern coast FL; Bahamas; Antilles; northern Gulf of Mexico 514-2063

C & B, 

2004b

Narella bellissima (Kukenthal, 1915)

Straits of FL (off Delray Beach); Bahamas; Lesser Antilles 161-792 C & B, 2003

Narella pauciflora Deichmann, 1936

Straits of FL (off Delray Beach); Bahamas; Cuba; Puerto Rico; Campeche Bank, Mexico 738-1473 C & B, 2003

Narella versluysi (Hickson, 1909)

Bermuda; Straits of FL (off St. Lucie Inlet, Palm Beach, Delray Beach;Bahamas); Cuba 677-900 C & B, 2003

Paracalyptrophora duplex Cairns & Bayer, 2004

Straits of FL (off Cape Canaveral - Cuba); Bahamas; Lesser Antilles 374-555

C & B, 

2004a

Paracalyptrophora simplex Cairns & Bayer, 2004

Insular side Straits of FL (off Palm Beach, north of Little Bahama Bank), Bahamas to Yucatan Channel 165-706

C & B, 

2004a

Plumarella aurea (Deichmann, 1936) off SC to Cuba 310-878

C & B, 

2004b

Plumarella dichotoma Cairns & Bayer, 2004 off SC to FL 494-1065

C & B, 

2004b

Plumarella laxiramosa Cairns & Bayer, 2004 off North and South Carolina 348-572

C & B, 

2004b
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STATE OF DEEP CORAL ECOSYSTEMS IN THE SOUTHEASTERN U.S. REGION

Higher Taxon Species Distribution Depth Range Reference

Plumarella pellucida Cairns & Bayer, 2004

off NC, through Straits of FL; Bahamas 549-1160

C & B, 

2004b

Plumarella pourtalesii (Verrill, 1883)

off NC, through Straits of FL; Cuba; Bahamas 183-882

C & B, 

2004b

Thouarella bipinnata Cairns, 2006

off northern FL; Straits of FL; off Little Bahama Bank; off Guyana 507-1000

Cairns, 

2006; 

Ross et al. 

unpub.

Order PennatulaceaFamily Kophobelemnidae Kophobelemnon sertum Verrill, 1885 off NC 1542 1 MRClass HydrozoaOrder AnthoathecataeSuborder FiliferaFamily Stylasteridae Crypthelia floridana Cairns,1986 eastern, southwestern FL 593-823

Cairns, 

1986

Distichopora foliacea Pourtalès, 1868

GA; Straits of FL (off FL Keys); SE Gulf of Mexico; Yucatan Channel (off Arrowsmith Bank) 183-527

Cairns, 

1986

Pliobothrus symmetricus Pourtalès, 1868 SC through Lesser Antilles

73-922; commonly 

150-400

Cairns, 

1986

Stylaster complanatus Pourtalès, 1867

GA; Bahamas; Yucatan Peninsula; Virgin Islands 183-707

Cairns, 

1986

Stylaster erubescens Pourtalès, 1868

SC - SW FL; Bahamas; Cay Sal Bank; Yucatan Channel (off Arrowsmith Bank)

146-965; 

commonly 650-

850

Cairns, 

1986

Stylaster laevigatus Cairns, 1986

SC; Bahamas; Cuba; Yucatan Channel (off Arrowsmith Bank)

123-759; 

commonly 300-

400

Cairns, 

1986

Stylaster miniatus (Pourtalès, 1868)

SC; Straits of FL (off FL Keys); Bahamas; Cuba 146-530

Cairns, 

1986
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271

STATE OF DEEP CORAL ECOSYSTEMS IN THE 

GULF OF MEXICO REGION: 

TEXAS TO THE FLORIDA STRAITS

Sandra Brooke1 and William W. Schroeder2

1

Ocean Research and Conservation Association, 

Fort Pierce, Florida 34949

2

Marine Science Program, University of Alabama, 

Dauphin Island Sea Lab, Dauphin Island, AL, 36528

I. INTRODUCTION

This report provides a summary of the current 

state of knowledge of deep (defined as >50 m) 

coral communities that occur on hard-bottom 

habitats in the Gulf of Mexico region, For the 

purposes of this report, the Gulf of Mexico region 

includes the waters within the U.S. exclusive 

economic zone (EEZ) of Texas, Louisiana, 

Mississippi, Alabama, and Florida as far north 

as Biscayne Bay on the East Coast (Figure 7.1), 

which includes the Pourtales Terrace and part 

of the Miami Terrace. The spatial distribution 

of deep coral species and their associated fauna 

are placed in the context of the geology and 

hydrography of three sub-regions, all of which 

have extensive deep coral communities, but 

with very different biological structure. The subregions are: (1) the northern Gulf of Mexico (2) 

the west Florida shelf and slope, and (3) the 

Florida Straits. The structure of the report follows 

the primary hydrographic flow from west to east 

through the Gulf of Mexico and into the Florida 

Straits. Threats affecting these communities, 

management and conservation concerns, and 

research needs are also discussed.

The most extensively documented coral habitats 

in the Gulf of Mexico are within the Flower 

Garden Banks National Marine Sanctuary 

(FGBNMS), which is located approximately 

160 km south of the Texas/Louisiana border. 

Since the inception of the sanctuary, most of 

the research has been focused on communities 

within SCUBA depths at the East and West 

Flower Garden Banks and Stetson Bank. Over 

the past few years however, investigation into 

the deeper habitats on the shelf, both within and 

outside the sanctuary boundaries, has expanded 

knowledge of the biological communities that 

are found below SCUBA depth limits. Extensive 

high-resolution multibeam mapping surveys have 

been conducted by NOAA, MMS and USGS, 

on select reefs and banks in the region (http://

walrus.wr.usgs.gov/pacmaps/wg-index.html), 

and this information has facilitated focused ROV 

and manned submersible operations.

The deep shelf and slope regions of the 

northern Gulf of Mexico have been extensively 

mapped and surveyed during exploration for oil 

and gas deposits, which led to the discovery 

and subsequent research on chemosynthetic 

communities associated with hydrocarbon 

seepage. The substrate in the deep Gulf of Mexico 

is principally comprised of fine particulates; 

however, large amounts of authigenic carbonate 

deposits are precipitated from biogeochemical 

activity associated with hydrocarbon fluid seepage 

(Schroeder 1992). Authigenic carbonates 

provide hard substrate for a wide variety of 

benthic fauna, including the structure-forming 

scleractinian, Lophelia pertusa. In 1955, Moore 

and Bullis collected large quantities of L. pertusa 

(=prolifera) in 420 to 512 m of water from the 

northeastern continental slope approximately 74 

km east of the Mississippi River delta (Moore and 

Bullis,1960). More recently, reports of living L. 

pertusa in the Gulf of Mexico are available from 

publications (Cairns 1979, 2000, Cairns and 

Viada 1987, McDonald et al. 1989, Schroeder 

2002, Schroeder et al. 2005), and records from the 

National Museum of Natural History Taxonomic 

Database. Recent research expeditions 

conducted between 2000-2004 (funded by 

NOAA’s office of Ocean Exploration and the 

Minerals Management Service) represent some 

of the first scientific submersible and remotely 

operated vehicle (ROV) dives in these areas, and 

they provide considerable new information on the 

distribution, habitat and biodiversity of deep coral 

communities in the Gulf of Mexico (Continental 

Shelf Associates, in review)
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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272

Figure 7.1 Map of the Gulf of Mexico and Florida, showing the Exclusive Economic Zone (EEZ) boundary. The 

Gulf of Mexico regional chapter covers deep water coral habitat enclosed by the red EEZ boundary and does 

not extend beyond the blue line. Map credit: Flower Garden Banks National Marine Sanctuary (FGBNMS)

Little is known about the ecology of the west 

Florida slope, although Collard and D’Asaro 

(1973) documented many benthic invertebrates 

of the eastern Gulf of Mexico from extensive 

dredging surveys, and Cairns (1978) compiled 

a comprehensive list of ahermatypic scleractinia 

for the entire Gulf of Mexico. Almost a decade 

later, Newton et al. (1987) described the coral 

mounds of the west Florida slope at depths 

of 500m. They found L. pertusa, Madrepora 

oculata and Bathypsammia sp, but none of 

their coral samples were living. More recent 

exploration of this area by Reed et al. (2004, 

2005b, 2006b) has expanded our descriptive 

knowledge of the fauna, but ecological questions 

remain unanswered.

Near the western end of the Straits of Florida, 

the Tortugas and Agassiz Valleys exhibit hardbottom habitats and high-relief escarpments at 

depths of 512-1,189 m (Minter et al. 1975). Deep, 

hard substrates may also exist in 500-1000 m 

depths on the Tortugas Terrace, 80 km west of 

the Dry Tortugas (Uchupi, 1968), but the fauna 

of these areas have not been explored. In the 

southern Straits of Florida and at the southern end 

of the Florida carbonate platform, the Pourtales 

Terrace provides extensive, high-relief, hard 

bottom habitat, at depths of 200-450 m. Louis de 

Pourtales discovered the feature in 1867 during 

a survey aboard the U.S. Coast Survey ship Bibb 

to lay a telegraph cable from Key West to Havana 

(Jordan et al., 1964). Alexander Agassiz (1888) 

named this feature the Pourtales Platform, and 

Jordan and Stewart (1961) later renamed it the 

Pourtales Terrace. Jordan (1964) discovered 

large sinkholes on the Pourtales Terrace. Land 

and Paull (2000) mapped and described nine of 

these sinkholes using side-scan sonar, seismic 

profiler, and echo-sounder profilers aboard the 

U.S. Navy’s submersible NR-1. Reed et al. 

(2005) also described the fish and invertebrate 

communities associated with high-relief deepwater structures and deep-water sinkholes on 

the Pourtales Terrace using the Johnson SeaLink submersible. 

II. GEOLOGICAL SETTING

The Gulf of Mexico basin consists of many different 

topographic features. The continental shelf 
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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273

slopes gradually to depths between 100 and 

200 m. The widest point is off southern Florida 

(about 300 km wide) and narrowest is at the 

Mississippi Delta (10 km). The continental shelf 

off west Florida and the Yucatan are carbonate 

with some complex topographic features; the 

eastern Gulf of Mexico and Texas-Louisiana 

shelves are primarily composed of terrigenous 

sediments. In the northern Gulf of Mexico, the 

Mississippi and Bryant submarine fans and the flat 

Sigsbee Abyssal Plain give way to the complex 

slump structure of the East Mexican Slope and 

the extremely complex topography of the Texas/

Louisiana continental slope (Rowe and Kennicutt 

2001). This section briefly describes the 

geology of each region to provide context for 

the more detailed description of their biological 

communities.

Northern Gulf of Mexico

The middle and outer portions of the Texas/

Louisiana shelf are scattered with intermittent 

banks of various depths and shape. These 

are usually comprised of carbonates, with silt 

and clay, overlying raised salt diaper structures 

(Rezak et al. 1985). These banks all support 

hard bottom communities, with different 

levels of complexity, but the most well known 

and ecologically well developed are the Flower 

Gardens Banks, which are situated on top of salt 

domes rising from approximately 130 m to 20 m 

depth (Figure 7.2). Salt domes began to form 

160-170 million years ago when salt layers 

were deposited in what was then a shallow sea, 

and subject to evaporation. In subsequent years, 

deep layers of sediments were deposited over 

the salt layers. Eventually, internal pressures 

became great enough to push isolated pockets 

of salt up through the sediments, forcing the 

seafloor to bulge upward in distinct domes. The 

Flower Gardens coral reef communities began 

developing on top of the domes 10,000 to 

15,000 years ago and have now overgrown the 

bedrock on which they developed.

The continental slope off Texas, Louisiana, 

and portions of Mississippi is geologically and 

physiographically one of the most complex in 

the world. Over ninety basins and seven 

submarine canyons dissect the continental 

margin of the northwestern Gulf of Mexico. Major 

portions of the middle and lower slope appear 

to be devoid of gas seeps, while the upper 

slope contains salt diapiric structures (such as 

mud and gas mounds), fluid expulsion features, 

hard-grounds, erosional gullies, numerous gas 

seeps and gas hydrate deposits. Deposits of 

authigenic carbonate, produced as a byproduct 

of chemosynthetic activity, provide hard substrate 

for development of sessile benthic communities, 

many of which are coral-dominated (Figures 7.3a, 

b). Eroded into the complex topography of the 

Texas/Louisiana slope, are four major canyon 

systems: the Mississippi, Keathley, Bryant, and 

Alaminos Canyon (Rowe and Kennicutt 2001). 

In the eastern portion of the region, sediments 

Figure 7.2. Map of the Flower Gardens area showing the location of the many reefs and banks in this area. 

Map credit: Resek et al. 1985.
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274

B

A Figure 7.3. Bathymetric map 

of the Gulf of Mexico 

showing known areas of 

abundant coral habitat. 

Chart from NOAA, NOS 

Bathymetric maps 1986. 

Scale 1:100,000. A) 

Viosca Knoll sites: dense

L. pertusa and other coral 

colonies at both VK826 and 

VK862/906. Map credit: 

OIMB for the U.S. Minerals Management Service 

(MMS).

B) Mississippi Canyon 

and Green Canyon sites: 

MC885 is comprised of 

large fields of C. americana 

delta and the Green Canyon 

sites support well developed 

L. pertusa. Map credit: 

OIMB for MMS.

from the Mississippi cover the western edge of the 

Florida shelf and a transition towards carbonate 

sediments begins. The Florida escarpment 

separates the Florida shelf from the Gulf Basin 

and also forms the southeastern side of the 

Desoto Canyon. In a region of high sediment 

deposition, the presence of the Desoto Canyon is 

poorly understood (Gore 1992). Some theories 

suggest that the canyon is the result of erosion 

caused by oceanic currents, possibly the Loop 

Current (Nowlin 1971).

West Florida shelf and slope

The west Florida shelf is a gently sloping (1-2o) 

broad carbonate platform that extends 750 km 

from Desoto Canyon in the north to the western 

Straits of Florida (Holmes, 1981). Along the 

edge of the west Florida shelf there is a series 

of “drowned reefs” or fossil reefs” at water depths 

ranging from approximately 50 m to over 120 m. 

In the northern region, small areas of the shelf 

have been designated as protected areas 

(Madison Swanson, Steamboat Lumps) by 
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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275

isobath on the southwest Florida slope is a 20 

km long zone of high-relief (10–15 m) Pleistocene 

coral mounds (Figure 7.4a). This region was 

first recognized from high resolution seismic 

reflection profiles during a cruise on the R/V 

Cape Hatteras in 1984 and consists of dozens 

and possibly hundreds of lithoherms (Figure 7.4b) 

composed of rugged black phosphorite-coated 

limestone boulders and outcrops (10- 15 m in 

height), some of which are capped with thickets 

of Lophelia coral (Newton et al. 1987, Reed et al. 

2004, Reed et al. 2006b). A deepwater sinkhole 

(200m) has also been described in detail from 

submersible dives off the west Florida shelf (Reed 

et al. 2005b). The Florida slope then grades 

into the Florida Escarpment, which extends 

from depths of 2500–3280 m into the eastern 

Gulf of Mexico. The face of the escarpment 

has steep vertical limestone cliffs of Cretaceous 

age, with intervening sediment-covered planes 

that provide habitat for dense chemosynthetic 

communities (Paull and Neumann 1987, Paull et 

al. 1990, 1991). 

Florida Straits

Two Miocene-age terraces, the Miami Terrace and 

Pourtales Terrace, occur off southeastern Florida 

and the Florida Keys reef tract. The Pourtales 

Terrace is a large triangular area over 213 km in 

length that runs parallel to the Florida Keys (Figure 

Figure 7.4a. Map of the 

eastern Gulf of Mexico 

showing the location of 

Lophelia lithoherms on the 

southwest Florida Slope 

(Reed et al. 2006). Scale 

1:100,000. Map credit: 

NOAA, NOS Bathymetric 

maps 1986. 

Figure 7.4b. Seabeam image of a southward view of the SW Florida Slope lithoherms (x2 vertical exaggeration). Image 

credit: Reed et al. and NOAA-OE.

the National Marine Fisheries Service (NMFS) 

on the recommendation of the Gulf of Mexico 

Fisheries Management Council (GMFMC) to 

preserve declining reef fish populations. on 

the southwest shelf edge, the shallowest of 

these ancient reefs, is Pulley Ridge, which 

is approximately 200 km long and supports 

well-developed communities of zooxanthellate 

scleractinian plate corals and other hard-bottom 

fauna at depths of 60-70 m. This area has been 

protected from bottom fishing to protect the fragile 

benthos, which is currently the deepest 

known hermatypic reef complex in U.S. Atlantic 

waters. Seaward of the shelf, at the 500m 
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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276

Figure 7.5. Chart of the Straits of Florida showing the Pourtales Terrace. The lithoherms are marked with black 

circles and the Jordan and Marathon sinkholes with black squares.(from Land and Paull 2000, Reed 2005b). 

Scale 1: 100,000. Map credit: NOAA, NOS Bathymetric maps 1986. 

7.5). The terrace is the drowned southern end of 

the Florida carbonate platform covering 3,429 km2

at depths of 200-450 m. The terrace provides 

extensive, high-relief, hard-bottom habitat with as 

much as 120 m vertical relief. The eastern section 

is comprised of a band of irregular topography and 

has the greatest relief on the terrace. The central 

section is flat with no topographic features. The 

southwest margin of the terrace contains a series 

of sinkholes that extend for approximately 100 km 

off the lower Florida Keys (Jordan and Stewart 

1961, Jordan et al. 1964, Land and Paull 2000). 

One of these, the Jordan Sinkhole, has a vertical 

relief of 206 m and may be one of the deepest 

and largest sinkholes known. The Jordan and 

Marathon sinkholes were described in 

detail from submersible dives by Reed et 

al. (2005b). The middle and eastern portion of 

the Pourtales Terrace consists of a northeasterly 

band of karst-like topography, with depressions, 

flanked by knolls and ridges that extend up to 

91 m above the terrace (Jordan et al. 1964, 

Land and Paull 2000). Further to the northeast 

is another zone of 40m high topographic relief 

that lacks any regular pattern (Gomberg 1976, 

Reed 2004). There are many high-relief bioherms 

(up to 120 m vertical relief) in this area, including 

a region called “The Humps” by local fishers, 

which is approximately 26 km south of Alligator 

and Tennessee Reefs in the Florida Keys (Reed 

2004, Reed et al. 2005b).

III. OCEANOGRAPHIC SETTING

The Gulf of Mexico is a semi-closed basin of 

approximately 1.5 million km2 with the continental 

shelves surrounding a deep abyss with maximum 

depths of approximately 3400 m in the eastern 

portion and 3700 m in the western portion. The 

surface waters of the Gulf of Mexico have been 

studied in great detail, but there is comparatively 

little information on circulation below 1000 m. 

Direct current measurements were rare (e. g., 

Pequegnat 1972, Hamilton 1990) until recently 

(Inoue et al. 2002, Hamilton and Lugo-Fernandez 

2001). The dominant hydrographic feature in 

the Gulf of Mexico is the Loop Current, which 

is formed when the Yucatan Current intrudes 

northward into the Gulf of Mexico from the 

Caribbean, flows clockwise around the basin, 

and empties into the Straits of Florida.
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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277

Northern Gulf of Mexico

While the Loop Current and associated rings 

are the major energetic currents in the Gulf of 

Mexico, creating extremely complex shelf and 

slope circulatory patterns, several other classes of 

energetic currents have also been observed in the 

deep waters of the Gulf (Bryant et al. 2000). Data 

collected in 1999 using the Texas A&M University 

(TAMU) deep-tow seismic system showed evidence 

of a large field of sedimentary furrows running 

along the base of the Sigsbee Escarpment in the 

Bryant Canyon area of the northwestern Gulf 

of Mexico (Bryant et al. 2000). Submarine 

furrows have been previously observed along 

the Blake Bahama Outer Ridge, the Bermuda 

Rise, the Brazilian Margin, and several other 

localities (see review by Flood, 1983), but the huge 

extent of the furrows observed in the northwestern 

Gulf of Mexico is unprecedented. Near-bottom 

current meters deployed near the base of the 

Sigsbee Escarpment (1978 m) have recorded 

flow events with velocities of more than 85 cm

s-1 at 2000m (Hamilton and Lugo-Fernandez, 

2001), during loop current eddy shedding. The 

effect of these strong deepwater flows on slope 

and shelf currents and their benthic communities 

is unknown and warrants further research.

West Florida Shelf and Slope

The Loop Current dominates the circulation in 

the eastern Gulf of Mexico and anticyclonic rings 

spawned by the Loop Current move west and 

south (Rowe and Kennicutt 2001). The Loop 

Current creates a vigorous north-south flow in 

the eastern Gulf of Mexico and as it migrates 

laterally over the west Florida shelf, it produces 

temperature variations at shallow to intermediate 

depths (50-500m), both seasonally and over 

longer time scales (Leipper et al. 1972).

Florida Straits

The Loop Current is joined in the Florida Straits 

by waters passing through the Old Bahama 

Channel to form the Florida Current, which in 

turn joins the Antilles Current in the Atlantic to 

form the Gulf Stream. Currents associated with 

the Loop Current can extend to great depths 

in the Florida Straits. High current speeds 

(>100 cm s-1) were observed at 200 m on the 

Pourtales Terrace (Reed at al. 2005b). Prior 

to this observation, lower speeds of 10 cm s-1 

had been recorded at 500 m depth (Cooper et 

al. 1990), but these persisted at one location for 

weeks to months. Strong current events have 

also been observed below 1000 m, suggesting 

that the Loop Current and eddies influence the 

hydrodynamics of the deepest portions of the 

Gulf of Mexico (Hamilton 1990).

IV. STRUCTURE AND HABITAT-FORMING

DEEP CORALS

The Gulf of Mexico has distinct regional 

differences among faunal communities (Table 

7.1). These include hermatypic corals in the 

shallower (<50 m) salt dome and carbonate shelf 

habitats, Lophelia-dominated communities on 

the hard substrate of the northern and eastern 

slope (>200m). Gorgonians and antipatharians 

are present to various degrees in all habitats 

but are primary structure-forming coral taxa in 

parts of the deep slope habitats in the northern 

Gulf of Mexico, on the shelf between 50-150 

m, associated with hardbottom features in and 

around the reefs and banks, and in parts of the 

deep slope habitats. Stylasterid hydrocorals 

are locally abundant on the Florida carbonate 

shelf and provide some structure but do not form 

large thickets. Three species of Pennatulacea 

(sea pens) have been recorded from the 

northern Gulf of Mexico (National Museum of 

Natural History (NMNH) database) but do not 

form significant structural habitat as in some other 

regions. There are also over 40 species of cup 

corals present throughout the Gulf of Mexico 

(Appendix 1), but again, these do not contribute 

greatly to habitat structure.

a. Stony corals (Class Anthozoa, Order

 Scleractinia)

The dominant azooxanthellate colonial 

scleractinian coral in the deep (>100m) Gulf of 

Mexico is Lophelia pertusa. Extensive thicket 

development occurs in the northern Gulf of 

Mexico, the southwest Florida lithoherms and 

on parts of the Pourtales and Miami Terraces 

(Schroeder 2002, Reed et al. 2005b, Reed 

et al. 2006). The most extensive L. pertusa 

habitat in the northern Gulf of Mexico is situated 

within the Minerals Management Service lease 

block Viosca Knoll 826. Further details are 

given below in the section on spatial distribution 

of coral species and habitats. The L. pertusa 

colonies found in the Gulf of Mexico exhibit two 

different morphologies; a heavily calcified thick 

branching structure which is seen at the Viosca 

Knoll area and a more fragile form with shorter 
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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278

internodes that is found elsewhere (Figure 7.6). 

An extensive study of L. pertusa habitats in the 

northern Gulf of Mexico, funded by the Minerals 

Management Service, was completed in 2005 

(Continental Shelf Associates in review). This study 

addressed various aspects of L. pertusa biology 

and ecology, including in situ growth of stained 

colonies, timing of gametogenesis, tolerance 

a range of sediment loads and temperatures, 

and community characterization. The staining 

experiment showed that L. pertusa growth was 

highly variable (some polyps grew and produced 

multiple new polyps, but others on the same 

branch did not grow at all) but on average the 

linear extension was between 2.4-3.36 mm yr-1. 

This is slightly lower than published growth data 

(e.g. average linear extension of 5.5 mm yr-1 by 

Mortensen and Rapp 1998), but was measured 

for the heavily calcified morphotype, which may 

be different from the more fragile growth forms. 

The gametogenic cycle of L. pertusa in the Gulf 

of Mexico is of similar duration to those in the 

Eastern Atlantic but is offset by several months. 

In the fjords of Norway, L. pertusa spawns in 

late February/early March and the subsequent 

gametogenic cycle has already begun before 

the prior cycle ends (Brooke and Jarnegren 

unpublished). In the Gulf of Mexico, mature 

oocytes were found in early September and 

primary oocytes in November, which indicates 

that spawning occurs sometime in October 

(Continental Shelf Associates in review). The 

causes of the difference in timing of reproduction 

are unknown and warrant further investigation. 

Temperature tolerance experiments showed that 

L. pertusa can survive for short periods of time 

at temperatures as high as 20o C for 24 hours, 

but long term survival requires a temperature 

between 10o C and 15o C. This corresponds to 

published observations that the upper thermal 

limit of L. pertusa distribution is approximately 

12o C (Rogers 1999). Sediment experiments 

conducted in the laboratory (Continental Shelf 

Associates in review) show that L. pertusa 

Table Key

Attribute Measure

Reef-Building Yes/No

Relative Abundance Low/ Medium/ High

Size (width or height) Small (<30cm)/ Medium (30cm-1m)/ Large (>1m)

Morphology Branching/ Non-branching

Associations None/ Few (1-2)/ Many (>2)

Spatial Dispersion Solitary/ Clumped

Overall Rating Low/ Medium/ High

Taxa

Reefbuilding AbundanceMaximum 

colony size Morphology

Associations 

with other 

structure forming invertebratesColony 

spatial 

dispersion

Overall rating 

of structural 

importance

Lophelia 

pertusa Yes Medium Large Branching Many

Solitary/ 

Clumped High

Madrepora 

oculata No Low Large Branching Few Solitary Medium

Callogorgia 

americana 

delta No Medium Large Branching Few/Many

Solitary/ 

Clumped Medium

Isididae No Medium

Medium/ 

Large Branching Few/Many Clumped Medium

Other 

Alcyonacea No Medium

Medium/ 

Large Branching Few/Many

Solitary/ 

Clumped Medium

Antipathidae No Medium Large Branching Few/Many

Solitary/ 

Clumped Medium

Stylasteridae No

High (Florida 

Straits) Small Branching Many

Solitary/ 

Clumped

High (Florida 

Straits)

Table 7.1. Structure-forming attributes of deep corals in the Gulf of Mexico region
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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279

could tolerate sediment loads of 54 mg L-1 for 

up to 2 weeks with approximately 90% survival, 

but increasing the sediment load to 103 mg L-1

caused almost 50% mortality over the same time 

period.

Other colonial species found in this region include 

Madrepora oculata (Linnaeus 1758), which forms 

large individual colonies on authigenic carbonate 

boulders in the northern Gulf of Mexico (Figure 7.7), 

but unlike L. pertusa, does not form monospecific 

coral stands and does not co-occur with L. pertusa 

thickets as habitat forming structure. Madrepora 

carolina has also been recorded from the 

northern Gulf of Mexico and the Florida Straits 

(Cairns 1979). In addition, Solenosmilia 

variabilis and Enallopsammia profunda are found 

on the carbonate slope habitats in the eastern 

Gulf of Mexico, but the extent of either species is 

unclear. 

In addition to these azooxanthellate scleractinian 

corals, there are areas in the Gulf of Mexico 

where zooxanthellate coral reefs have been found 

at depths greater than 50m. These areas include 

the reefs of the East and West Flower Gardens 

Banks and the unique communities at McGrail 

Bank, which are dominated by Stephanocoenia sp. 

(Schmahl and Hickerson 2006). Zooxanthellate 

structure-forming scleractinians are also found 

on Pulley Ridge on the southwest Florida shelf, 

where zooxanthellate corals such as Agaricia

sp., thrive at depths of 60-70m. Since these 

are not true deepwater corals, but rather shallow 

water species that have extended into deeper 

water, they will not be addressed in detail in this 

report.

b. Black corals (Class Anthozoa, Order

 Antipatharia)

At least 20 species of antipatharians have 

been documented for the Gulf of Mexico 

region (Appendix 1) with at least half of these 

identified from the Flower Gardens Banks 

area. Other samples have been collected and 

are awaiting identification or description (S. 

Brooke pers. obs.). Black corals are locally very 

common in some areas of the northern Gulf of 

Mexico and the Florida Straits. Some species 

grow into large bushy colonies (eg Leiopathes 

Figure 7.6. Samples of Lophelia pertusa colonies collected from the Gulf of Mexico showing (left 

morph) heavily calcified morphology (brachycephala) with large polyps from VK826, and (right 

morph) fragile morphology (gracilis) from GC354. Photo credit: S. Brooke OIMB.
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sp.) which may provide substrate and refuge for 

other organisms. Unlike some of the other taxa, 

there seems to limited geographical separation in 

the distribution of most of the species. Exceptions 

to this observation are the Leiopathes species, 

which were only documented for the northern 

Gulf of Mexico. Three colony colors (red, orange 

and white) have been collected and deposited 

at the National Museum of Natural History, but 

it is still unclear whether they are conspecific 

color morphs or distinct species (D. Opresko 

pers.comm.). The most thorough investigation on 

reproduction of black corals was conducted 

on Antipathes fiordensis from the southwestern 

New Zealand Fjords (Parker et al. 1997). Colonies 

were gonochoristic, and gametogenesis was rapid 

and synchronous, beginning in November and 

terminating in March. Colonies reached sexual 

maturity between the heights of 70 and 105 cm 

which corresponded to a minimum age for sexual 

maturity of about 31 yr. Such information is not 

available for the deep Gulf of Mexico species 

therefore it is unknown whether late reproductive 

maturity is a characteristic of this taxa.

c. Gold corals (Class Anthozoa, Order

 Zoanthidea)

Gold corals are not known in this region.

d. Gorgonians (Class Anthozoa, Order

 Alcyonacea)

There are numerous species of octocorals in 

the deep waters of the Gulf of Mexico (many of 

which are still unidentified), the majority of which 

belong to the family Plexauridae; however Cairns 

and Bayer (2002) have identified several species 

of the habitat-forming primnoid Callogorgia spp. 

occurring throughout the Gulf of Mexico. At least 

one subspecies, Callogorgia americana delta is 

endemic. Most of these species contribute to 

multi-species coral habitats, but some occur as 

dense monotypic assemblages in localized areas 

in the northern Gulf of Mexico. These include 

the bamboo coral Acanella arbuscula (family 

Isididae) and C. americana delta, which is known 

to provide nursery habitat for oviparous cat sharks 

that deposit their egg cases on the branches 

(Etnoyer and Warrenchuck in press, S. Brooke 

pers. obs.), and is often seen with large fleshy 

ophiuroids (Asteroschema sp.) entangled in the 

colonies (Figure 7.8). A list of octocorals found 

in the Gulf of Mexico region can be found 

in Appendix 1. A comprehensive octocoral 

species inventory is currently being updated by 

Dr. S. Cairns of the National Museum of Natural 

History for publication in 2007. Gorgonians are 

an important component of the shelf edge reefs 

and banks of the northwestern Gulf of Mexico 

Figure 7.7. Large 

colony of Madrepora 

oculata showing 

distinctive zig-zag 

branch morphology. 

Photo credit: Brooke et 

al. and NOAA-OE
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between 50 and 120m depth, and the FGBNMS 

research team is developing catalogs of 

octocorals, antipatharians and sponges found 

on these features (E. Hickerson pers.comm.). 

Despite being an important component of many 

deepwater hard-bottom communities, octocoral 

biology is not well understood. A study of 

reproduction in multiple species of common 

deepwater octocorals is currently underway 

(Simpson, pers.comm.). Information to date 

shows that there is variation in reproductive 

strategy within this order (Fitzsimmons–Sosa et 

al. 2004, Brooke unpublished data).

e. True soft corals (Class Anthozoa, Order

 Alcyonacea)

Several species of soft corals have been 

documented for this region (Appendix 1). 

Anthomastus (Bathyalcyon) robustus delta was 

often found in hardbottom habitats of the northern 

Gulf of Mexico at ~274m and at shallower depths 

(50-135m) Chironepthya (=Siphonogorgia) 

caribaea was encountered regularly in surveys 

of hardbottom areas in the NW GOM (E. 

Hickerson pers. comm.). None of the soft 

coral species contribute significantly to habitat 

structure since they neither form large colonies 

nor have solid skeletons. Very little is known 

about the biology of these taxa.

f. Pennatulaceans (Class Anthozoa, Order

 Pennatulacea)

There are at least two species of pennatulaceans 

recorded for the Gulf of Mexico, and these were 

collected at greater depths (2683m) than the 

majority of the other cnidaria in the region (NMNH 

database). These taxa are often found on soft 

substrate and are the dominant benthic fauna in 

some areas (e.g. Halipteris willemoisi in Alaska); 

however there is insufficient information on 

distribution and abundance to determine whether 

this is the case for the pennatulaceans in the Gulf 

of Mexico.

g. Stylasterids (Class Hydrozoa, Order

 Anthoathecatae, family Stylasteridae)

Stylasterid hydrocorals are common components 

of the benthic communities of the bioherms and 

sinkholes of the Florida Straits, and the SW Florida 

shelf lithoherms, but have not been recorded in 

Figure 7.8. Callogorgia americana delta, a gorgonian from the northern Gulf of Mexico, showing Euryalid 

ophiuroids entwined in the branches. Photo credit: OIMB for MMS.
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the northern Gulf of Mexico basin. Stylasterids are 

usually absent from areas with high suspendedsediment, which may be detrimental to both 

juvenile and adult colonies (Ostarello 1973, 

Cairns 1992). They generally have slow growth 

rates, long life, and brooded larvae, and may be 

out-competed by more aggressive species in high 

nutrient waters, finding refuge in low nutrient 

and/or less competitive environments such as 

cryptic and deep-water habitats (Sanders 1979, 

Thayer 1989, Cairns 1992). A list of stylasterid 

species currently documented for the Gulf of 

Mexico, is presented in Appendix 7.1. Taxonomic 

descriptors follow revisions of the taxa by Cairns 

(1986). Dense communities of stylasterids 

have been observed on the bioherms of the 

central and eastern Pourtales Terrace, primarily 

along the tops and flanks of the structures 

(Reed et al. 2005b) and thick piles of dead 

and live stylasterid colonies were observed in 

some locations, but without coring the interior, 

it was not possible to ascertain how much 

contribution these stylasterid colonies made to 

the accumulation of the bioherms. Stylasterids 

are also one of the dominant sessile taxa of 

the southwest Florida Shelf Lophelia lithoherms 

(Reed 2004, Reed et al. 2006b)

V. SPATIAL DISTRIBUTION OF CORAL

SPECIES AND HABITATS

The distribution of coral communities in 

the Gulf of Mexico region can be divided by 

geologic setting and depth. Dozens of reefs and 

banks are scattered across the shelf edge of the 

Gulf of Mexico, particularly in the northwestern 

region. Many of these features are formed by salt 

diaparism, which have pushed the sediments up 

into the photic zone. On the shallowest of these 

domes, zooxanthellate coral reefs and coral 

communities have developed, and are flanked by 

diverse communities of octocorals, antipatharians, 

and sponges. Authigenic carbonate deposits 

Figure 7.9. An example of deep coral habitat at the Flower Garden Banks National Marine Sanctuary, typical 

of the NW Gulf of Mexico habitats. Image includes octocorals, antipatharians, echinoderms, sponges, octocorals and deep water fishes. Photo credit FGNMS/NURC-UNCW.
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along the deeper parts of the northern shelf 

and slope provide hard substrate for L. pertusa 

communities as well as other scleractinians 

octocorals and antipatharians (see Cairns et al. 

1994). Zooxanthellate coral reefs are also found 

along the top of the west Florida shelf (<100m), 

and, at deeper depths, the west Florida slope and 

the terraces of the Florida Straits support extensive 

L. pertusa formations as well as stylasterine and 

sponge-dominated communities (Reed et al. 

2005b).

Northern Gulf of Mexico

The benthos of the northwestern Gulf of 

Mexico is composed of terrigenous sediments, 

interspersed with dozens of reefs and banks, 

many of which are connected by corridors of 

hardbottom features. The East and West Flower 

Garden Banks, within the FGBNMS, support 

the northernmost zooxanthellate coral reefs in 

the U.S.A., with scleractinian coral cover of up 

to 70%, which is much higher than comparable 

coral reefs in the Caribbean basin (Precht et 

al. 2005; Rezak et al. 1985). These banks are 

just two of many reefs and banks that support 

rich communities of antipatharians, octocorals 

and sponges at depths between 50 and 150m 

(Figure 7.9). The FGBNMS has developed a 

database of information derived from more than 

8300 digital images, taken during >160 ROV 

surveys in the northwestern GOM. In addition, 

approximately 200 antipatharian, octocoral 

and sponge samples were collected in order to 

produce regional catalogs for each taxon. 

The deeper regions on the northern Gulf of Mexico 

slope cannot support zooxanthellate corals but 

are known to have fairly extensive areas of L. 

pertusa communities. These are located in an 

area known as Green Canyon off Louisiana and 

on the upper flanks and on Viosca Knoll, a deepwater salt dome off Alabama and Mississippi. 

More detailed distribution patterns are described 

below.

Viosca Knoll

The most well developed and well-documented 

L. pertusa communities in the Gulf of Mexico 

occur on the southwest flank of a mound in the 

southwest corner of Viosca Knoll lease block 826 

(29o 09.5′ N 88o 01.0′ W, 430-520 m) on the upper 

DeSoto slope. Bottom sediments consist of 

authigenic carbonate and unconsolidated clay, 

silty clay, disarticulated shells, and shell hash. 

Authigenic carbonate formations are abundant 

at this site, especially on the crest and flanks of 

the mound, and occur in the form of large plates, 

slabs, and irregular shaped blocks, boulders, 

and rubble (Schroeder 2002). Representative 

ranges of near-bottom temperature, salinity, and 

dissolved oxygen values for the coral habitat 

obtained ~13 km east of the site), are 7.0-9.3o 

C, 34.9-35.1 psu, and 2.6-3.2 ml L-1,respectively 

(Schroeder 2002).

Hard substrate on the crest and flanks of the 

mound support large and abundant colonies of 

L. pertusa (Schroeder 2002). These colonies 

have a bushy morphology composed of irregular, 

dendritic branches that are highly anastomosed 

and heavily calcified with large polyps. Individual 

colonies range in size from a few centimeters to 

over 1.5 m in diameter, while aggregations of 

closely associated colonies attain 1.5 to 2 m in 

height and width and 3 to 4m in length. Many of 

the aggregated colonies appear to be in the first 

phase of the “thicket” building stage described 

by Squires (1964). Colonies less than 25 to 50 

cm in diameter were predominantly 100% live. 

Larger colonies and aggregated colonies often 

had dead branches (light to dark brown in 

color) at their base and center with live terminal 

branches, and some were 100% dead coral. This 

site is associated with a mature chemosynthethic 

community, consisting primarily of living 

tubeworm aggregations, but no known mussel 

beds or hydrates. 

In 1955, the M/V Oregon collected approximately 

136 kg of L. pertusa from a deep-water reef 

system, comprised of several sections up to 300 

m long and 55 m deep (Moore and Bullis, 1960). 

The coordinates of the sampling site (29o 05.0’N, 

88o 19.0′W, 421-512m) were revisited by the U.S. 

Navy submarine NR-1 in 2002 (Schroeder pers. 

obs.), but there were no L. pertusa reefs found 

at this location. The depth recorder tracing 

presented by Moore and Bullis (1960) is similar to 

the cross-sectional profile of a submarine canyon 

in lease block VK 862/906 (29o 06.4′N, 88o 

22.9’W). This area is comprised of a topographic 

high located on the northern edge of an exposed 

carbonate rock complex that extends southward 

for over 2 km to the eastern rim of the canyon. 

Water depths range between 300 - 500 m. 

This lease block region has been explored using 

ROV (Sonsub Innovator), submersible (JSL) 

and submarine (NR-1) but the L. pertusa habitat 
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284

described by Moore and Bullis has not been 

located. This reef may have been buried by 

a slumping event, or simply has not yet been 

found.

An unidentified species (or several species) 

of white anemone dominates the fauna in this 

region (Figure 7.10). They can be extremely 

abundant (up to 94 m-2) and occur on flat 

substrate as well as carbonate outcroppings 

(Figure 7.11), and often co-occur with an 

unidentified species of hexactinellid sponge. 

The large carbonate boulders support diverse 

assemblages of corals including L. pertusa 

colonies, dense aggregations of bamboo corals 

(Isididae), C. americana delta (Primnoidae) and 

large black corals (Antipathidiae) (Continental 

Shelf Associates in review)

Mississippi Canyon

A low relief mound-like sea floor feature with 

abundant authigenic carbonate rock lies within 

Mississippi Canyon lease block 885 at a depth 

of 625 m (28o 03.78’N, 89o 42.62’W). It is an area 

of active seepage and includes bacterial mats, 

mussel beds, shells of dead clams, tubeworm 

aggregations and brine seepage. Interspersed 

among these areas of active seepage are 

carbonate outcrops with small Lophelia 

pertusa colonies and larger carbonate 

mounds with larger colonies of both L. 

pertusa and Madrepora oculata. There 

are also areas of high C. americana delta 

abundance, but overall coral species 

diversity at this site is low (Continental 

Shelf Associates in review)

Green Canyon

Thickets of aggregated L. pertusa 

colonies exist along a single ridge in 

an area of Green Canyon lease block 

234 (27o 44. 81 ’N, 91o 13.13 ’W). The 

ridge is approximately 130 m in length 

and is comprised predominantly of 

dead Lophelia pertusa with live outer 

branches. Some of the thickets are 

large (several m3) with occasional 

colonies of 100% live coral. On one 

side of this ridge is a large area with 

extremely abundant Callogorgia 

americana delta colonies. This is an 

area of active seepage, as evidenced 

by tubeworm aggregations observed 

close to the coral ridge.

Green Canyon lease block 184/185, commonly 

known as Bush Hill, is a low-relief knoll located 

in approximately 580 m of water. In addition 

to numerous aggregations of old tubeworms, 

both gas hydrates and mussel beds are present 

at this site. The central chemosynthetic 

communities are bordered on the northwest 

corner of the knoll by a series of large authigenic 

carbonate outcrops. Abundant large gorgonians 

(C. americana delta and other species) and 

colonies of L. pertusa are present on these 

outcrops and on an escarpment on the western 

side of the knoll.

The explored part of Green Canyon lease 

block 354 (between 27o 35.9’N 91o 49.6’W 

and 27o 35.8’N, 91o 49.4’W) is part of a slope 

that descends from 520 to 560 m. Abundant 

authigenic carbonate boulders on the upper 

portion of the slope support large Lophelia 

pertusa thickets of 5 to 10 m in diameter. 

Scattered vestimentiferans and some large 

tubeworm aggregations are interspersed with 

these boulders. Down-slope of this area, there 

are occasional large pockmarks with slumping 

sediments and authigenic carbonate outcrops 

supporting smaller L. pertusa and Madrepora 

Figure 7.10. Image of unidentified white anemones and hexactinellid sponges found in high numbers at VK862 (Figure 7.3) 

Photo credit: OIMB for MMS.
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oculata colonies (Continental Shelf Associates 

in review)

.

West Florida Shelf and Slope

There are numerous hardbottom habitats along 

the west Florida shelf from Panama City to 

the Dry Tortugas (Schroeder et al. 1989). One 

of the most well-known is the Florida Middle 

Grounds in the eastern Gulf of Mexico. The 

bank formations consist of two parallel ridges 

separated by a valley and 23 species of stony 

coral and 170 species of fish have been recorded 

from this area (Nipper et al. 2006). Other high 

relief reefs off the western Florida panhandle 

are associated with the rim of the De Soto 

Canyon. These reefs are popular fishing grounds 

for snapper and grouper and include Madison 

Swanson, Mud Banks and Twin Ridges to the 

northeast and lower relief structure such as The 

Edges and Steamboat Lumps. Southern shelf 

edge reef areas include Howel Hook, Hambone 

Ridge, Northwest Peaks, Christmas Ridge and 

Pulley Ridge (Koenig et al. 2000b). Pulley Ridge 

is a series of drowned, barrier islands on the 

southwest Florida Shelf which form a ridge about 

5 km across with less than 10 m relief. The 

shallowest parts of the ridge are about 60 m deep 

and the southern portion supports zooxanthellate 

scleractinian corals, making it the deepest 

hermatypic reef in the U.S.A. Atlantic. The corals 

Agaricia sp. and Leptoceris cucullata are most 

abundant, forming plates up to 50 cm in diameter. 

Less common species include Montastraea 

cavernosa, Madracis formosa, Madracis decactis, 

Porites divaricata, and Oculina tellena (Halley et 

al. 2005). Deepwater ahermatypic coral mounds 

occur along the 500 m isobath of the west Florida 

Slope for approximately 20 km between 26o 20’N, 

84o 45’W to 26o 30N, 84o 50’W, with individual coral 

mounds between 5 and 15 m tall. The lithoherms 

consist of rugged black phosphorite-coated 

limestone boulders and outcrops capped with 

0.5-1.0 m tall thickets of L. pertusa (Reed et al. 

2006). The R/V Aleutian Bounty collected the 

first recorded samples of Lophelia pertusa and 

Madrepora oculata from this area (26o30.0′ N, 

84o50.0′ W, 640 m) in a trawl net in 1983. In 1984 

the R/V Cape Hatteras also collected samples 

of corals from the mounds using rock dredges 

(Newton et al. 1987). Colonies of M. oculata 

were also reported at much lower abundance, 

together with a solitary coral Bathypsammia 

Figure 7.11. This image shows the different kinds of anemones that represent the dominant benthic fauna at VK862 (See figure 7.3). Photos credit: OIMB for MMS.
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sp. All corals recovered in the 1984 collections 

were dead; however in 2003, the SONSUB 

ROV was used to ground-truth three of the slope 

features: a 36-m tall escarpment and two of the 

many lithoherms (Reed et al. 2004, Reed and 

Wright 2004, Reed et al. 2005b). The dominant 

fauna on the 36 m escarpment consisted of an 

antipatharian species (approx 30 cm high), an 

isidid bamboo coral (30-40 cm high) and numerous 

other octocorals, plus several species of sponge 

(Heterotella sp., Phakellia sp., Corallistidae). The 

benthic communities of the lithoherms differed 

from those of the escarpment in that thickets of 

live and dead L. pertusa were found on some 

of the slope terraces and the top ridges. Coral 

cover was estimated at <5% to >50% in some 

areas, but was only 1-20% alive (Reed et al. 

2006). Dominant sessile macrofauna included 

stony corals, octocorals, stylasterid hydrocorals, 

black corals (Antipathes sp. and Cirrhipathes sp.) 

and sponges from the families Hexactinellidae 

and Demospongiae (Reed et al. 2004).

Florida Straits

Between 1999 and 2005, the Clelia and 

Johnson-Sea-Link (JSL) submersibles were 

used to survey several high relief sites on 

the Pourtales Terrace, including the Jordan 

and Marathon sinkholes and five of the high 

relief lithoherms on the central portion of the 

terrace (Reed et al. 2004, 2005b). The peaks 

of some of the mounds were covered with thick 

layers of stylasterid corals along with dense 

and diverse communities of sponges and 

octocorals (Reed et al., 2005b). Cnidarians 

included 3 species of antipatharian black 

coral, 5 stylasterid hydrocorals, 11 octocorals, 

and 1 scleractinian. Stylasterid and antipatharian 

corals were also common on the flat pavement 

adjacent to the base of the mounds. High 

densities of sponges, stylasterid corals and 

octocorals were observed, particularly on the 

bioherm plateaus and terraces. Sponges and 

stylasterids also dominated slopes of the 

bioherms but at much lower densities than the 

plateaus, whereas the octocorals were generally 

found at higher densities on the slopes (Reed et 

al. 2005b). Certain species occurred only on the 

Pourtales sinkholes and were not found on the 

bioherms; these included two identified species 

of antipatharians (Antipathes rigida and A. 

tanacetum), three species of octocorals 

(Paramuricea placomus, Plumarella pourtalesii 

and Trachimuricea hirta) and the scleractinian 

Solenosmilia variabilis.

Along the eastern edge of the Miami Terrace 

at a depth of 365 m is a 90 m tall steep rock 

ridge capped with L. pertusa, stylasterid corals, 

bamboo coral, black coral, and various sponges 

and octocorals (Reed et al. 2004, 2006b). The 

benthic communities of the Pourtales Terrace 

bioherms however, differ from those of the 

lithoherms along the northeastern Straits of 

Florida (Messing et al. 1990) primarily due to 

an absence of L. pertusa and a dominance of 

stylasterid corals and stalked crinoids (Reed et 

al. 2004, 2006b).

Shipwrecks

There are 7,000 reported shipwrecks in the Gulf 

of Mexico. While most pose no threat to the 

environment many were carrying cargoes of 

fuel and other materials and these may lose their 

structural integrity over time. The contribution of 

these wrecks to habitat development is currently 

unknown; they may serve as temporary 

substrate for coral colonization and structure 

for fish and invertebrate populations, until their 

deterioration releases fuel and other pollutants. 

They also may serve as an initial focus for 

development of an established habitat.

The ‘Deep Wrecks Project’ funded by the 

Minerals Management Service and NOAA, 

with researchers from C&C Technologies, 

Inc., used an industrial ROV (Sonsub Inc) to 

archaeologically and ecologically assess seven 

World War II era vessels sunk by U-boats in 

the Gulf of Mexico. Since these sites ranged in 

depths from 87 m to almost 2,000 m, they offered 

biologists a unique opportunity to study the 

artificial reef effect in differing ecological niches. 

Three of the vessels, the Virginia, Halo and 

Gulfpenn, are located south of the Mississippi 

delta in water depths of 87 m, 143 m and 554 

m, respectively and supported several deep 

sea coral families. Scleractinians were found 

on all three wrecks, with four of the five species 

occurring on the Virginia (Madracis myriaster, 

Oculina varicosa, Paracyathus pulchellus and 

Pourtalosmilia conferta), two species on the Halo 

(M. myriaster and P. conferta) and two species 

on the Gulfpenn (P. conferta and L. pertusa). 

Gorgonians were collected from and/or 

observed on the two shallowest wrecks. The 

Halo had a well-developed gorgonian fauna, 

including four species that were collected from 
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287

the wreck (Placogorgia rudis, Thesea sp. cf. T. 

grandiflora, Thesea sp. cf. T. rubra and Thesea 

sp.) and one unidentified species recorded on 

video. A single, large colony of Muricea pendula 

was found on the Virginia, together with two 

species of antipatharians (Antipathes furcata and 

Stichopathes sp. cf. S. pourtalesi), which occurred 

only on the Virginia. All species, except Oculina 

varicosa, have previously been reported from the 

Gulf of Mexico.

VI. SPECIES ASSOCIATIONS WITH DEEP

CORAL COMMUNITIES

Information on deepwater coral-associated 

fauna in the Gulf of Mexico region comes 

from historic trawl and dredge samples (e.g., 

Agassiz 1869, Moore and Bullis 1960, Newton 

et al. 1987), and also from recent ROV and 

submersible operations.

Northern Gulf of Mexico

Antipatharian, octocoral, and sponge 

communities associated with hardbottom 

areas in the Northwestern Gulf of Mexico 

contain abundant populations of both 

commercial and non-commercial fish species, 

including Lutjanus campechanus (red snapper),

Rhomboplites aurorubens (vermillion snapper), 

Pronotogrammus martinicensis (roughtongue 

bass), Anthias tenuis (threadnose bass), and 

several species of grouper. Initial observations 

from the FGBNMS research suggest correlations 

between the benthic community structure and 

composition and the mobile fauna (E. Hickerson 

pers. comm.). Rezak et al. (1985) provide a 

list of fish associated with reefs and banks in the 

northwestern Gulf of Mexico.

Unlike many other Lophelia reefs, the northern 

Gulf of Mexico L. pertusa communities have not 

been shown to support commercially important 

fisheries species. Large schools of Hyperoglyphe 

perciformis (barrelfish) have been observed at 

Viosca Knoll lease block 862 (S. Brooke pers. 

obs.). These are caught along with Polyprion 

americanus (wreckfish) along the U.S. eastern 

coast, and they are marketed along with Beryx 

splendens and B. decadactylus (alfonsinos). 

Although there is currently no commercial fishery 

for barrelfish in the northern Gulf of Mexico, there 

is potential for development of a future fishery for 

this species. Information on Lophelia-associated 

fish species has been documented (Sulak et al, 

in press) during a series of cruises in the northern 

Gulf of Mexico. 

Commercially important deepwater 

invertebrates that exist in the northern Gulf 

include Pleoticus robustus (royal red shrimp)

and Chaceon fenneri (golden crab) (Snyder 2000), 

although there is currently no fishery for golden 

crab species in the northern Gulf of Mexico. 

Royal red shrimp occur over specific substrata in 

different areas: terrigenous silt and silty clay off 

the Mississippi Delta and calcareous mud off the 

Dry Tortugas, with peak abundance at 250-500 m 

depth. Golden crabs occur over a similar depth 

range, but prefer hard bottom and outcroppings 

such as the Florida escarpment (Lindberg and 

Lockhart 1993). The Moore and Bullis site (located 

in 1955) discussed previously, was reported to 

be ‘over half a mile long and up to 180 ft thick’. 

Records indicate a single trawl drag recovered 

“several hundred pounds of fish, shrimp, starfish, 

urchins and other animals”. This site has not 

been found since that time, but recent research 

in the nearby Viosca Knoll area (Continental 

Shelf Associates in review) has shown that 

L. pertusa habitat supports a similar coralassociated community as that reported by Moore 

and Bullis. In addition, L. pertusa colonies provide 

habitat for the surrounding slope community, but 

probably do not directly provide food to other 

organisms, with the exception of the gastropod 

Coralliophila sp. Some species are found 

at higher densities close to L. pertusa or have 

been found only in tight association with coral 

habitat. These species include the squat lobster 

Eumunida picta, comatulid crinoids, unidentified 

sponge species and, in dead coral skeleton only, 

sabellid polychates (Continental Shelf Associates 

in review).

West Florida Shelf and Slope

NMFS landing statistics show higher landings of 

deepwater reef species than the rest of the Gulf. 

For the grouper family (Serranidae), the highest 

catches were for Epinephelus flavolimbatus

(yellowedge grouper), which are associated 

with low-relief hard bottom habitats up to 250 m 

deep. Other serranid species such as E. niveatus

(snowy grouper), E. drummondhayi (speckled 

hind) and E. nigritus (warsaw grouper) were also 

found on deepwater hard bottom structure, but 

landings of these species were consistently much 

lower than for E. flavolimbatus.. Hard-bottom 
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288

species such as Etelis oculatus (queen 

snapper) and Hyperoglyphe perciformis

(barrelfish) were minor contributors to the 

commercial fishery. These data encompass 

the entire west coast of Florida, including the 

popular shallower fishing grounds on the Florida 

shelf and shelf edge. These habitats serve as 

spawning aggregation sites for commercially 

valuable species such as Mycteroperca 

microlepis and M. phenax (gag and scamp)

as well as habitat for other commercial grouper 

and snapper species. In June 2000 the Madison 

Swanson/Steamboat Lumps marine reserves 

were implemented (with a 4 year sunset clause 

that was extended in 2004), and all fishing was 

banned except trolling for highly migratory 

and coastal pelagic species between May 1st 

and October 31st. This was done in an effort 

to protect grouper spawning stocks and has 

proved a successful policy, despite illegal fishing, 

which is still a problem on these offshore areas 

(Koenig pers. comm.). There are no landings 

data currently available specifically for the 

deepwater coral lithoherms, and since many 

are deeper than 250 m, the shallower nearshore 

regions are probably contributing the majority 

of the catch. There is also a small fishery for 

Chaceon fenneri (golden crab), which reached 

a peak of 640 metric tons in 1995, but by 2004 

had declined to 25 metric tons. During a 

study of golden crab distribution and behavior 

(Lindberg and Lockhart 1993), the greatest 

density was found among gorgonians and in 

crevices on hard bottom at depths of 350-550 

m. The coordinates of the survey correspond 

to the lithoherm region on the west Florida slope, 

but fishing data do not indicate precisely where 

landed crabs were caught.

Dominant sessile macrofauna on the west 

Florida slope lithoherms included stony corals, 

gorgonians, stylasterids, antipatharians and 

sponges (Reed and Wright, 2004). These coral 

habitats provide complex structure for abundant 

and diverse communities of associated fish, 

crustaceans (including the golden crab, C. 

fenneri), mollusks, echinoderms, polychaete 

and sipunculan worms, anemones, terebratulid 

brachiopods, bryozoans and other sessile and 

motile macrofauna, some of which are probably 

undescribed species (Newton et al. 1987, 

Reed 2004). Preliminary studies discovered 

new species of octocorals and sponges from 

some of these sites (Reed et al. 2004). Messing 

(unpublished) found an unstalked crinoid 

assemblage (159 m) not known to occur 

elsewhere in Florida but characteristic of deeper 

Cuban and Bahamian waters (>300 m).

Florida Straits

A total of 31 fish taxa (of which 24 were identified 

to species level) have been identified from the 

sinkhole and bioherm sites of the Pourtales 

Terrace (Reed et al. 2004). Common species on 

the high-relief bioherms included Antigonia capros

(deepbody boarfish), Lopholatilus microps (blueline 

tilefish), Epinephelus niveatus (snowy grouper), 

and Holanthias martinensis (roughtongue bass). 

Some species, such as the snowy grouper, 

blackbelly rosefish (Helicolenus dactylopterus) 

and mora (Laemonema melanurum), were 

common at both the sinkhole and bioherm sites. 

This region is shallower in places than the SW 

Florida lithoherms or the Northern Gulf of Mexico 

regions, and supports commercially valuable 

species such as E. drummondhayi (speckled 

hind), E. flavolimbatus (yellowedge grouper), E. 

nigritus (warsaw grouper), Pagrus pagrus (red 

porgy), Seriola sp. (amberjack) and Pareques 

iwamotoi (blackbar drum), as well as deeper 

species such as E. niveatus (snowy grouper), H. 

dactylopterus (blackbelly rosefish), Lopholatilus 

sp. (tilefish) and Urophycis sp. (phycid hakes). 

The fish densities appeared insufficient to support 

a viable commercial or recreational fishery, since 

only a few individuals of the larger grouper 

species were present at any one site (Reed 

2004). A swordfish was also observed on top 

of Pourtales Terrace from the NR-1 submersible 

(C. Paull, pers. obs.), and the JSL submersible 

was attacked twice during dives on the Terrace 

and sinkholes (Reed pers.comm.) 

There is extremely limited information on coral 

associated fauna from the Pourtales Terrace. 

Trawl collections from the Terrace (Agassiz 

1869) contained an abundant and unique 

benthic assemblage of invertebrates rarely found 

elsewhere in the Straits of Florida, e.g., the 

hydroid Cladocarpus sigma, and the unstalked 

crinoids Comatonia cristata and Coccometra 

hageni (Bogle 1975, Meyer et al. 1978). The most 

recent published information on the communities 

of the Florida straits deepwater habitats are 

by Reed (2004) and Reed et al. (2005b). These 

manuscripts document the coral communities 

and their associated fauna collected during 

submersible and ROV cruises, but represent a 
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small percentage of the highly biodiverse habitat 

assemblages.

VII. STRESSORS ON DEEP CORAL

COMMUNITIES

The most significant global threats to deep corals are 

destructive fishing practices, particularly trawling, 

that compromises the habitat infrastructure, and 

often occurs repeatedly in the same area, thus 

preventing and habitat recovery; however, trawling 

is not a significant problem in the deep Gulf of 

Mexico (Table 7.2). Another potential stressor 

is the expanding exploration and exploitation of 

fossil fuel reserves in the deep Gulf of Mexico.

Fishery Effects

Bottom trawling

Trawling for the various varieties of shrimp 

(Farfantepenaeus duorarum, Penaeus setiferus, 

P. aztecus) represents the most lucrative fishery 

in the Gulf of Mexico; however these shrimp are 

fished over soft sediments at less than 50 m depth 

therefore it is unlikely that these shrimp fisheries 

have damaged deep coral habitats. Trawling 

for Pleoticus robustus (royal red shrimp) occurs 

in water depths of 400-500 m off Florida Texas 

and Alabama. Gear used for royal red shrimp is 

similar to trawling gear for shallow species, but 

to work in deepwater the trawl doors and lines 

must be much heavier. Since royal red shrimp 

inhabits soft substrate (silt and calcareous mud) 

this fishery probably does not target deepwater 

coral habitat, but damage could occur if the 

shrimp habitat is adjacent to coral communities 

or if equipment is erroneously deployed in coral 

habitat. There are currently no other deepwater 

trawl fisheries operating in this region.

Scallop dredges

There is a dredge fishery in the Gulf of Mexico 

for Argopecten gibbus (calico scallops); it is 

infrequent but when scallops are present in high 

numbers, vessels come from as far as the midAtlantic coast to participate in the fishery (Arnold 

2006; NMFS 1982), and there is some potential 

for impacts to deep coral since these scallops 

can be found in water depths of 10-400 m. The 

present condition of stock in the South Atlantic 

region is unknown because of large fluctuations 

in abundance and the last commercial landings 

of calico scallops from the Gulf coast of Florida 

was in 2000 (http://research.myfwc.com) 

Deep Gill Nets

Gillnets are allowed for the spanish and king 

mackerel fishery, and for non-FMP commercial 

and recreational fisheries. There is no deep water 

managed species that is fished using gillnets.

Bottom Long-lines

Fishing for ‘highly migratory species’ such as 

tuna and swordfish (Xiphias gladius) is one of 

the most lucrative in the Gulf of Mexico, primarily 

using surface long-lines. These are not deep 

enough to interfere with coral habitat, but damage 

could occur however if the lines break and drag 

across the coral habitat. Many sharks are fished 

with bottom longlines, particularly on the shelfslope break and these have potential to damage 

fragile benthos.

Gear Type

Potential 

Severity of 

Impact

Potential Extent 

of Impact from 

Fishing Gear

Current 

Geographic 

Extent of Use in 

Region

Overlap of 

use with coral 

habitat

Overall 

Rating 

of Gear 

Impact

Bottom Trawl High High Low Uncertain Low-Medium

Mid-water Trawl Low Low Medium Uncertain Uncertain

Dredge High Low N/A N/A N/A

Bottom-set Longline Medium Low Low Uncertain Low

Bottom-set Gillnet Medium Medium N/A N/A N/A

Longline traps or pots High Medium Low Uncertain Uncertain

Single pots Low Medium Low Uncertain Uncertain

Hook and line Low Low Low to medium* Yes Low

*may increase as recreational trophy fishing increases

Table 7.2 Potential and Current Fishing Gear Impacts on Deep Sea Corals
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Demersal or reef fishes caught in deepwater 

include Epinephelus flavolimbatus (yellowedge 

grouper), E. niveatus (snowy grouper), E. 

nigritus (warsaw grouper), Etelis oculatus (queen 

snapper) and Lutjanus vivanus (silk snapper), 

plus Caulolatilus microps (blueline tilefish) and 

Caulolatilus chrysops (goldface tilefish), which 

are generally found on soft sediment benthos 

rather than reefs or hardbottom. These 

species are primarily harvested using bottom 

long-lines, with some traps or hook and lines 

used. Bottom long-lines are much shorter than 

pelagic lines, but can be up to 30 km long (Snyder 

2000). These lines can damage fragile coral 

habitat, especially in areas of high current 

where lines may be dragged or heavily weighted. 

Numerous lost long-lines have been observed 

on the seafloor during submersible dives in the 

deepwater Naples sinkhole (200m) on the west 

Florida shelf, and deepwater sinkholes and 

bioherms of the Pourtales Terrrace (Reed pers.

comm.) 

Traps and Pots

Golden crab gear consists of rectangular wire 

mesh traps that are attached in series along a 

weighted mainline and are recovered by hauling 

the mainline aboard the boat. Golden crabs have 

been observed on hard bottom habitat in the 

region of the Florida lithoherms (Lindberg and 

Lockhart 1993) and are fished on the Pourtales 

Terrace (Reed pers.comm.). If the traps were 

placed close to the coral, recovery of the trap 

could result in dragging it over the coral and 

potentially damaging the coral.

Harvest of precious corals

There is currently no commercial harvesting of 

precious corals in this region; however there 

are areas that support abundant populations 

of antipatharians, which are harvested in other 

areas (e.g. Hawaii) for making ornaments and 

jewelry.

Effects of Other Human Activities

Oil and Gas Exploration and Extraction

The Gulf of Mexico supports the largest 

offshore oil and gas extraction fields in the 

nation. Currently, there are approximately 4,000 

producing platforms, of which about 1,962 are 

major platforms and approximately 152 companies 

are active in the Gulf of Mexico (http://www.gomr.

mms.gov) (Figure 7.12). The deep waters 

of the Gulf of Mexico (>300 m) where deep 

coral resources may occur have been an area of 

increasing exploration and development over the 

past decade. Deep-water oil production has risen 

386% since 1996 and accounts for 62% of the 

total oil production in the Gulf of Mexico. By 

Figure 7.12. Active oil and gas leases in the Gulf of Mexico. Map credit: MMS
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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291

the end of 1999, approximately 1,200 wells had 

been drilled in water depths greater than 1000 

ft (305 m). In 2003, an estimated 350 million 

barrels were produced from deepwater rigs. 

Deep water gas production has increased by 407% 

since 1996, reaching an estimated 1.42 trillion 

cubic feet (Tcf) in 2003, and has surpassed 

shallow water production (U.S. Department of 

the Interior 2000, 2004).

Direct physical impact to the substrate has 

been observed at Viosca Knoll 826 (Schroeder 

2002). At 496 m on the western flank, a furrow 

was observed in the soft sediment that extended 

both up- and down-slope. The furrow alternated 

between a completely disrupted surface over a 1-

1.5 m swath, and a clean, narrow cut into the soft 

sediment. There was no extensive destruction 

of carbonate structures; however, numerous 

colonies and aggregations of L. pertusa, attached 

to buried carbonate material, were severely 

damaged. It was surmised that this disruption 

occurred when a wire anchor cable, deployed in 

conjunction with oil and gas drilling operations 

in this region, struck the bottom one or more 

times. Apart from the physical impact, the other 

primary environmental threat from oil and gas 

exploration/extraction is accidental leakage of 

drilling fluids or oil into the water column. Drilling 

in deep water requires special drilling fluids that 

can operate at low temperatures. Syntheticbased drill fluids (SBF) have been used in the 

Gulf of Mexico since 1992 and are well suited 

to deepwater use (Boland et al. 2004). 

These fluids do not contain the toxic polycyclic 

aromatic hydrocarbons that are found in nonsynthetic drill fluids that are refined from crude 

oil. Use of SBF in the Gulf of Mexico is limited to 

rapidly-biodegradable esters and internal olefins; 

however, the accidental release of drilling fluid 

still presents potential environmental hazards 

through release of large quantities of sediment 

and anaerobic decomposition of the organic SBF 

material at the sediment surface. Drilling fluid 

may also be mixed at the time of release with oil 

containing toxic components.

The number of chemical spills in the Gulf of 

Mexico has steadily increased. In 1998 27% of 

all incidents in U.S. waters took place in the Gulf 

of Mexico (Boehm et al. 2001). In 2003, an SBF 

spill occurred when a drilling pipe fractured in two 

places at an oil lease block site in Mississippi 

Canyon (MC778), located at 28.19°N, 88.49°W 

at a depth of 1840 m. An environmental impact 

assessment conducted by MMS (Boland et al. 

2004) concluded that the SBF dispersed into the 

water, settled to the seafloor and biodegraded. 

The SBF would cause a temporary decrease in 

dissolved oxygen at the sediment water interface 

therefore the less motile animals within the 

nearby benthic community (mud bottom with low 

biodiversity) could have smothered under a layer 

of SBF or from anoxic conditions resulting from 

biodegradation, but that the community would 

recover once the SBF biodegrades. In this case, 

there were no strong currents, the sediment 

settled close to the accident and fortuitously did not 

impact a sensitive habitat, but this may not always 

be the case. If oil rather than SBF was released, 

the fluid would be more toxic, although it may be 

carried vertically away from the benthos since it is 

less dense than water. Interestingly, the report 

mentions that the closest chemosynthetic 

communities were at VK826, and were not close 

enough to be impacted by the spill, but made 

no mention of potential impact to the extensive 

Lophelia communities in the same lease block. 

Deposition of organic and potentially toxic 

sediment could cause extensive coral mortality 

from direct toxicity, smothering by sediments, 

and reduction in oxygen content. These reefs 

are slow growing and their recruitment rates are 

unknown. As oil and gas exploration increases, 

it is vital that corals are included in the MMS 

environmental impact assessments and are 

given the same status as chemosynthetic 

communities.

Deployment of Gas Pipelines and 

Communication Cables

The deployment of gas pipelines and cables 

can impact deep coral communities through 

sediment re-suspension during the burial process 

(especially in shallower areas), and can cause 

habitat destruction during pipeline deployment 

as a direct result of the pipes and also from 

anchor-cable sweep and anchor drag. While 

there has been a decline in the production of 

natural gas from the shallower existing lease areas 

in the Gulf of Mexico, the Minerals Management 

Service (MMS) began offering incentives in 2001 to 

encourage drilling in deeper waters. Koenig et 

al. (2000a) reported on the potential impact of 

gas pipelines proposed for deployment in two 

marine protected areas in the Gulf of Mexico 

and evaluated the potential destructive effects 

of the process. MMS regulations do not require 
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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292

burial and anchoring of pipelines laid at depths 

greater than 61 m. Other habitat impacts may 

be associated with deployment of pipelines. 

Each anchor measures approximately 5 x 6 m 

and weighs at least 13 tons and together with the 

cables cover a swath approximately one nautical 

mile wide As they are positioning the anchors, the 

cables act like trawls or dredges as they sweep 

across the bottom. The combined effects of the 

anchor-cable sweep, anchor drag and sea state 

determines the extent of the habitat damage.

The extent of the damage caused by pipeline 

construction is evaluated by MMS using clearly 

defined habitat descriptions. The MMS considers 

live-bottom to be “seagrass communities or those 

areas which contain biological assemblages 

consisting of sessile invertebrates, such as sea 

fans, sea whips, hydroids, anemones, ascidians, 

sponges, bryozoans, or corals living upon and 

attached to naturally-occurring hard or rocky 

formations with rough, broken, or smooth 

topography; or areas whose lithotope favors the 

accumulation of turtles, fishes, and other fauna” 

(Koenig et al. 2000a). Since the MMS does 

not require oil and gas exploration companies 

to survey areas at a depth greater than 100 

m important habitats including coral areas may 

be exposed to potential damage or destruction. 

These habitats include: essential fish habitat 

(EFH) for Lopholatilus chamaeleonticeps

(tilefish), Caulolatilus microps (blueline 

tilefish), Epinephelus nigritus (warsaw grouper), 

E. niveatus (snowy grouper), E. drummondhayi 

(speckled hind), and E. flavolimbatus (yellowedge 

grouper) (Parker and Mays 1998). The southwest 

Florida shelf has been found to have extensive 

live-bottom coverage (65%) at depths of 120 to 

160 m (Phillips et al. 1990), however the number, 

extent and location of deep coral habitats in the 

Gulf of Mexico are unknown.

Gulf Fiber Corporation installed a deep water fiber 

optic cable system called Fiber Web in 1999. 

This has been updated over the past 3 years 

and stretches from Texas to Florida in a loop that 

runs from onshore to the 200 m isobath, along 

the shelf edge and back onshore again. These 

cables are not thick and heavy like pipelines, 

but there is potential for environmental impact 

when they are laid or replaced. In 2003, Florida 

Department of Environmental Protection initiated 

a policy that directs underwater communications 

cables to be sited through gaps in reef tracts. This 

protects reefs under Florida state jurisdiction, but 

there is no such legislation for deep water corals 

in federal waters.

Mineral mining

The Gulf of Mexico has extremely large 

stores of methane hydrate, which is a solid 

crystal lattice comprised of water and methane. 

Research over the past two decades has shown 

that methane hydrate exists as void-filling in 

shallow sediments, and as massive mounds, 

often associated with chemosynthetic biota. At 

the moment, the extraction of these hydrates 

is in the experimental phase, but there is an 

interagency road map in place for the research and 

development of methane hydrate as a potential 

future energy source (U.S. Department of Energy, 

2006). In the deep Gulf of Mexico, corals are often 

found close to chemosynthetic activity, since the 

hard substrate is of authigenic origin. Methane 

hydrate extraction is not an immediate threat but 

could have future consequences for deep water 

chemosynthetic and coral communities.

Sedimentation

There are natural events that could temporarily 

increase sedimentation, such as benthic storms 

or deepwater gyre activity, but unless they are 

unusually severe or protracted, the sessile benthic 

fauna will have evolved to deal with such events. 

Anthropogenic sediment input could be caused 

by fossil fuel exploration or extraction, not only 

by resuspension of local sediments but also from 

drilling muds. Release of petroleum products or 

other extraction related contaminants is also a 

potential threat to benthic organisms. Currently 

the MMS Notice to Lessees issued in 2000 states 

that “If you propose activities that could disturb 

seafloor areas in water depths 400 meters 

(1,312 feet) or greater, maintain the following 

separation distances from features or areas 

that could support high-density chemosynthetic 

communities: 1) At least 1,500 feet from each 

proposed muds and cuttings discharge location 

and 2) At least 250 feet from the location of all 

other proposed seafloor disturbances (including 

those caused by anchors, anchor chains, wire 

ropes, seafloor template installation, and pipeline 

construction).” This notice does not mention 

avoidance of coral communities.
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VIII. MANAGEMENT OF FISHERY

RESOURCES AND HABITATS

Fisheries in the area covered by this report 

are managed undr fishery management plans 

(FMPs) by two regional councils, the Gulf of 

Mexico Fishery Management Council (GMFMC) 

and the South Atlantic Fishery Management 

Council (SAFMC). The Gulf of Mexico Fishery 

Management Council is responsible for fisheries 

in federal waters off the coasts from Texas to the 

west coast of Florida (www.gulfcouncil.org) and 

the South Atlantic Fishery Management Council 

is responsible for fisheries in federal waters off 

the coasts of North Carolina, South Carolina, 

Georgia and east Florida to Key West (http://

www.safmc.net).

The GMFMC produced a Coral and Coral Reefs 

FMP with the SAFMC in 1982. The species 

currently covered under this FMP include “species 

belonging to the Orders Stolonifera, Telestacea, 

Alcyonacea (soft corals), Gorgonacea (horny 

corals, sea fans, sea whips), and Pennatulacea 

(sea pens) in the Subclass Octocorallia; Orders 

Scleractinia (stony corals) and Antipatharia 

(black corals) in the Subclass Zoantharia; and 

the Orders Milleporina (fire corals, stinging 

corals) and Stylasterina in the Class Hydrozoa”. 

These classes include many deep coral species 

and therefore apply to deep corals as well 

as their shallow water counterparts. In March 

2005, the GMFMC proposed amending their 

description of coral EFH to “the total distribution 

of coral species and life stages throughout the 

Gulf of Mexico including: coral reefs in the North 

and South Tortugas Ecological Reserves, East 

and West Flower Garden Banks, McGrail Bank, 

and the southern portion of Pulley Ridge; hard 

bottom areas scattered along the pinnacles, reefs, 

and banks from Texas to Mississippi along the 

shelf edge and at the Florida Middle Grounds; 

the southwest tip of the Florida reef tract, and 

the predominant patchy hard bottom offshore 

of Florida from approximately Crystal River 

south to the Keys”. 

On January 23, 2006, the National Marine 

Fisheries Service established Habitat Areas 

of Particular Concern proposed by the Gulf of 

Mexico Fisheries Management Council. Fishing 

restrictions 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. 

Additional restrictions on fishing were already in 

place at the Tortugas Ecological Reserves and 

Madison-Swanson and Steamboat Lumps Marine 

Reserves. Other Gulf of Mexico HAPC’s that do 

not carry any regulations are the remainder of 

Pulley Ridge, the Florida Middle Grounds, and 

the following banks: 29 Fathom, MacNeil, Rezak, 

Sidner, Rankin, Bright, Geyer, Bouma, Sonnier, 

Alderdice, and Jakkula Banks. No regulations 

are currently in place for these HAPC areas, but 

they will be considered during individual fishery 

plan review and development.

These EFH HAPC were targeted toward the 

zooxanthellate coral communities, some of 

which occur at deeper depths than usual, but 

include areas that are biologically significant 

due to their antipatharian, octocoral, and sponge 

communities. The HAPCs do not include habitats 

where the structure-forming azooxanthellate 

scleractinians such as L. pertusa are found. 

Currently there are no proposed protected areas 

for these communities which are only found at 

depths >300 m in the Gulf of Mexico. The SAFMC 

however, has proposed the designation of HAPC 

status to several areas within their jurisdiction, 

including the deep-water coral habitats on the 

Pourtales Terrace. For further information and 

updates, refer to the SAFMC website. 

Flower Gardens Banks National Marine 

Sanctuary

In the 1970’s researchers and recreational divers 

initiated a 20 year effort to protect the reefs of 

the Flower Gardens Banks, culminating in the 

designation of the FGBNMS in 1992, which 

comprised the Eastern and Western Flower 

Gardens Banks. In October, 1996, Congress 

expanded the Sanctuary by adding Stetson 

Bank, which is a small salt dome located about 

110 km south of Galveston, Texas. The total area 

of the sanctuary encompasses 144 km2, which 

includes 1.6 km2 of reef crest at approximately 

18-25m, with zooxanthellate corals occuring to 

at least 52 m. Fishing within the Sanctuary is 

regulated through fishery management plans 

developed in cooperation with the Gulf of Mexico 

Fishery Management Council. In addition to the 

original protected areas, HAPC designations 

were granted in 2006 for selected deeper banks 

noted above. These sites were proposed to the 

GMFMC by the FGBNMS based on the deepwater 
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communities of octocorals, gorgonians, and 

antipatharians.

IX. REGIONAL PRIORITIES TO

UNDERSTAND AND CONSERVE DEEP 

CORAL COMMUNITIES

Deepwater corals exist throughout the Gulf of 

Mexico however their distribution has not been 

comprehensively surveyed or mapped and 

there are a very limited number of publications 

on coral habitat or associated fauna (Agassiz 

1869, Moore and Bullis 1960, Bogle 1975, 

Meyer et al. 1978, Newton et al. 1987, Lindberg 

and Lockhart 1993, Schroeder 2002, Reed 2004, 

Reed et al. 2004, Reed and Wright 2004, Reed 

et al. 2006). Most of the information regarding 

deep coral habitat is either anecdotal, proprietary 

or in preparation from current projects. NOAA’s 

Office of Ocean Exploration has supported, and 

continues to support, exploration of deepwater 

coral habitat throughout the U.S. EEZ. The 

Flower Garden Banks NMS approaches 

deepwater investigations on a regional basis, and 

will continue to look for opportunities to further 

the science in the northwestern Gulf of Mexico. 

The Minerals Management Service has recently 

funded a 3-year research project to characterize 

coral habitat, associated fauna and coral biology 

in the northern Gulf of Mexico (Continental Shelf 

Associates in review), but there is still a need 

for additional research in many areas, some of 

which are listed below.

Map coral habitat

• Deep coral habitat may be far more 

extensive than current knowledge indicates. 

In shallower areas on the shelf (<200m), 

there are extensive areas of hardbottom, 

produced primarily through salt tectonics, 

which provide substrate for coral community 

development. The deeper slope and basins 

are dominated by soft sediment habitats, 

but sessile organisms (such as corals and 

sponges) colonize exposed rocky outcrops 

of authigenic carbonate. The geology of the 

Gulf of Mexico has been well studied, primarily 

because of fossil fuel interests, so there is 

some indication of the distribution of different 

substrate types. This information has been 

used (and should continue to be used) as a 

starting point for ‘ground-truthing’ explorations. 

Areas of special ecological importance should 

be defined so that protection can be proactive 

rather than reactionary.

Priority mapping areas

• The high numbers of hard bottom 

lithoherms on the Southwest Florida slope, 

together with the diverse habitats discovered in 

the few explorations undertaken in this region 

indicate tremendous potential for unexplored 

coral and fish habitat.

• The Pourtales Terrace region has a 

different community composition from the 

other two regions and the limited exploration 

to date has shown community difference 

between sinkholes and lithoherms, which 

raise interesting ecological questions. Also, 

this region supports several potential fisheries 

species, but little is known of their abundance 

or ecological association with the corals.

• In the Northwestern Gulf of Mexico, the 

Flower Gardens NMS has identified a priority 

for mapping geological features between 

banks that may connect them, biologically 

and ecologically. Limited groundtruthing has 

revealed deepwater assemblages including 

octocorals, antipatharians and sponges 

associated with these features.

Identify and describe areas of EFH

• Since the GMFMC manages deep coral 

species under its Coral and Coral Reefs FMP 

the protection measures available under 

EFH regulations should provide a framework 

for the application of the Magnuson-Stevens 

Fishery Conservation and Management 

Act (National Marine Fisheries Service, 

1996). Most areas currently identified by 

the Council primarily reflect the distribution 

of shallow-water hermatypic corals. There 

is a need to identify the locations of deep 

coral communities, particularly in those areas 

that may be subject to future trawling (e.g., 

for royal red shrimp) or other anthropogenic 

impacts (e.g., pipeline construction or other 

oil and gas-related activities).

Increase research of important but understudied 

taxonomic groups

• Most deepwater coral research to 

date has focused on the structure-forming 

scleractinia; however, members of the 

antipatharia, gorgonacea and stylasterids 

also provide habitat structure and may be an 

important part of an ecosystem. Taxonomic 
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

GULF OF MEXICO

295

and some distributional data are available 

from museum inventories and institutional 

databases; however, with the increase in 

deepwater exploration in recent years, the 

databases may not be current. There are 

also many images available from underwater 

still and video cameras, but in many cases 

these cannot provide definitive identification. 

Additionally almost nothing is known about the 

ecology of these important groups.

• The porifera are also in urgent need of 

study, but the identification of the many taxa 

that contribute to habitat complexity is an 

extensive undertaking.

With the increasing level of oil and gas exploration 

in the northern Gulf of Mexico, surveying for 

fossil fuel exploration and environmental impact 

assessments by the energy industry could 

potentially provide video footage and anecdotal 

information on coral distribution. These data are 

not always available; however with increased 

cooperation between industry and science, such 

information could become a valuable resource. 

The SERPENT (Scientific and Environmental ROV 

Partnership using Existing Industrial Technology) 

program is an example of successful cooperation 

between multiple industry and scientific partners 

around the globe, including the Gulf of Mexico 

(http://www.serpentproject.com).

Cooperation with the MMS could potentially 

change environmental impact and protection 

policies to include areas of diverse hard bottom 

communities in the deep Gulf of Mexico and 

elsewhere. Policies could be implemented to 

protect coral communities and EFH from the 

direct effects of exploration and extraction of oil 

and gas, including deployment of cables and 

pipelines. Additionally, buffer zones could be 

created so that fossil fuel operations would be 

excluded within a ‘reasonable’ distance (to be 

defined) from coral habitat, to minimize the effect 

of construction and accidents.

X. CONCLUSION

The extent of deepwater coral habitat in the Gulf of 

Mexico is unknown at present, but in general, where 

there is hard substrate there is some degree of 

benthic community cover, and this is frequently 

dominated by cnidarians. Coral communities 

are found under various environmental 

conditions, from the hard substrate scattered 

throughout the shelf and slope of the northern 

Gulf of Mexico, to the carbonate shelf and slope 

of the west Florida shelf and the Florida Straits. 

Corals require hard substrate for settlement and 

sufficient water flow to deliver food and oxygen, 

and remove waste products, but not so much 

that feeding mechanisms or skeletal growth is 

impacted. They require particulate food since 

the true deepwater corals are azooxanthellate 

and must obtain their nutrition from zooplankton 

(possibly augmented by marine detritus 

and dissolved organic material), therefore 

factors that drive zooplankton dynamics may 

also influence coral distribution. Sediment load 

may affect coral communities, particularly those 

groups with very small polyps such as the 

stylasterids. The removal of fine particles from 

coral polyps is facilitated by mucous shedding 

and requires energetic expenditure. Finally (but 

not exhaustively) currents must deliver larvae to 

the appropriate habitats. There are large areas of 

bare, apparently suitable substrate in the northern 

Gulf of Mexico that have not been colonized (S. 

Brooke pers. obs.). Recruitment limitation is one 

possible explanation, but our understanding of the 

ecology of these deepwater coral habitats is still 

rudimentary.

The role of L. pertusa in structuring the 

surrounding slope community appears to be 

through the provision of habitat rather than direct 

provision of nutrition by L. pertusa, with the possible 

exception of the gastropod Coralliophila sp. L. 

pertusa creates habitat for a number of associated 

species, many of which show significantly higher 

densities in proximity to coral or have been 

found only in tight association with L. pertusa 

in the Gulf of Mexico. These include Eumunida 

picta (a species of squat lobster commonly seen 

on deepwater coral habitats in the southern 

U.S.A.), comatulid crinoids, sabellid polychaetes, 

and sponges (Continental Shelf Associates, in 

review). The coral associated species found 

in the Gulf of Mexico appear similar to those 

encountered on the L. pertusa reefs off the east 

coast of Florida, although there has not yet been 

a rigorous comparison. The coral communities in 

the Florida Straits are different from those of the 

northern Gulf of Mexico and the west Florida shelf 

and are dominated by stylasterine and sponge 

communities. Several species of Pleurotomarid 

gastropods are found on the shallower carbonate 

slopes and sinkholes on the Pourtales Terrace, 
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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but are not found in the Gulf of Mexico basin. 

Of all the deepwater coral communities addressed 

in this report, the only ones that show evidence of 

human impact are the shallower fishing grounds 

on the west Florida shelf. The grouper spawning 

aggregations on these mounds have been 

severely overfished (Koenig et al. 2000b), which 

prompted the GMFMC to grant protected status 

to Madison Swanson and Steamboat Lumps. 

The reefs and banks on the continental shelf 

edge of the northern Gulf of Mexico are protected 

through various regulatory agencies (NOAA’s 

National Marine Fisheries Service and Minerals 

Management Service). Regulations do not 

overlap, which may allow environmental impacts to 

occur, such as dynamiting coral reefs for salvage 

purposes, and methods of closing these gaps 

should be investigated and proposed. Whatever 

these alternatives are, should also be considered for 

off-shelf deepwater communities. The golden crab 

fishery could potentially damage coral on the west 

Florida slope, especially if the fishery expands in the 

future, but there are currently no major commercial 

trawl fisheries in the Gulf of Mexico to seriously 

impact deepwater coral habitat. The oil and gas 

industry is expanding rapidly in deepwater and 

will probably continue to do so far into the future. 

The laying of pipelines and cables, boat anchors 

and spills could all potentially cause extensive 

damage to coral habitat unless activity is restricted 

by regulations and enforced appropriately. The 

most well studied and most extensive L. pertusa 

habitat in the Gulf of Mexico is within the the 

Viosca Knoll lease block 826, and the adjacent 

lease block 862 also supports a highly biodiverse 

area with abundant fish populations (Continental 

Shelf Associates in review). Although there is 

no apparent immediate threat, these areas 

are unique as far as current investigation can 

determine and as such are worthy of protection 

from potential anthropogenic impact. Another 

region of interest is the lithoherm habitat on the 

west Florida shelf edge. The extent and health 

of this system is unknown and is a high priority for 

further research. There are many areas of the 

deep Gulf of Mexico that have not been explored 

and further investigation may discover more coral 

habitats. At present there are fewer immediate 

threats to Gulf of Mexico ecosystems than in 

certain other areas, but a proactive approach to 

habitat protection may be more effective than 

one which reacts to user groups that are already 

invested in the exploitation of a resource.

XI. REFERENCES

Agassiz L (1869) Report upon deepsea dredgings 

in the Gulfstream during the 3rd cruise

of the US steamer Bibb. Bulletin of the 

Museum of Comparative Zoology at 

Harvard University 1:363-386

Agassiz A (1888) Three cruises of the Blake. 

Houghton, New York 

Bogle MA (1975) A review and preliminary 

revision of the Agalaopheniiae (Hydroida: 

Plumulariidae) of the tropical western 

Atlantic. M.Sc thesis, University of 

Miami, 307pp

Boehm P, Turton D, Raval A, Caudle D, French 

D, Rabalais N, Spies R, Johnson J (2001) 

Deepwater Program: Literature Review, 

Environmental Risks of Chemical Products 

Used in Gulf of Mexico Deepwater Oil 

and Gas Operations; Volume I: Technical 

Report. OCS Study MMS 2001-011. 

Boland GS, Current C, Gravois M, Metcalf M, 

Peuluer E (2004) Fate and Effects of a 

Spill of Synthetic-Based Drilling Fluid 

at Mississippi Canyon Block 778. U.S. 

Department of the Interior, Minerals 

Management Service, Gulf of Mexico 

OCS Region report number OCS 2004-

039 

Bryant WR, Dellapenna T, Silva A, Dunlap W, 

Bean D (2000) Mega-furrows on the 

continental rise south of the Sigsbee 

Escarpment, AAPG Annual Meeting: New 

Orleans, LA (available online at www.

aapg.org)

Cairns SD (1978) A checklist of the ahermatypic 

scleractinia of the Gulf of Mexico,

with a description of a new species. Gulf 

Research Reports 6:9-15

Cairns SD (1979) The deepwater scleractinia of 

the Caribbean Sea and adjacent waters. 

Stud Fauna Curacao 50, p341 

Cairns SD (l986) A revision of the Northwest 

Atlantic Stylasteridae (Coelenterata: 

Hydrozoa). Smithsonian Contributions to 

Zoology 4l8: l3l pp
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

GULF OF MEXICO

297

Cairns SD (1992) Worldwide distribution of 

the Stylasteridae (Cnidaria: Hydrozoa). 

Scientia Marina. (Barc.) 56:2-3

Cairns SD (2000) A revision of the shallowwater azooxanthellate Scleractinia of the 

Western Atlantic. Studies Natural History 

of the Caribbean Region 75:1-231

Cairns SD, Bayer FM (2002) Studies on Western 

Atlantic Octocorallia (Coelenterata: 

anthozoa). Part 2: the genus Callogorgia 

Gray, 1858. Proceedings of the Biological 

Society of Washington 115: 840-867

Cairns SD, Opresko DM, Hopkins TS, Schroeder 

WW (1994) New records of deep-water 

Cnidaria (Scleractinia & Antipatharia) 

from the Gulf of Mexico. Northeast Gulf 

Science 13: 1-11

Cairns SD, Viada ST (1987) Range extensions 

of ahermatypic scleractinia in the Gulf of 

Mexico. Northeast Gulf Science 9:31-134

Collard SB, D’Asaro CN (1973) Benthic 

Invertebrates of the Eastern Gulf of Mexico. 

IN: Jones JL, Ring RE, Rinkel MO and Smith 

RE Eds. A summary of knowledge of the 

Eastern Gulf of Mexico. State University 

of Florida Institute of Oceanography, St 

Petersburg, FL, ppIIIG1-27

Continental Shelf Associates (in review) 

Characterization of Northern Gulf of Mexico 

deepwater hard bottom communities with 

emphasis on Lophelia coral. Submitted 

as final report to the U.S. Dept. of the 

Interior, Minerals Management Service, 

Gulf of Mexico OCS Region

Cooper C, Forristall GS, Joyce TM (1990) Velocity 

and hydrographic structure of two Gulf 

of Mexico warm-core rings. Journal of 

Geophysical Research 95:1663-1679

Etnoyer P, Warrenchuk J (in press) A catshark 

nursery in a deep gorgonian field in 

Mississippi Canyon, Gulf of Mexico. 

Bulletin of Marine Science

Fitzsimmons-Sosa K, Hallock P, Wheaton J, 

Hackett KE, Callahan MK (2004) Annual 

Cycles of Gonadal Development of Six 

Common Gorgonians from Biscayne 

National Park, Florida, USA. Caribbean 

Journal of Science 40:144-150

Flood R. D (1983) Classification of sedimentary 

furrows and a model for furrow initiation and 

evolution: Geological Society of America 

Bulletin 94:630-639

Gomberg D (1976) Geology of the Pourtalès 

Terrace, Straits of Florida. Ph.D. 

Thesis, Univ. Miami, Florida, p371 

Gore RH (1992) The Gulf of Mexico. Pineapple 

Press, Inc. Sarasota, Florida

Halley R, Dennis GP, Weaver D, Coleman F 

(2005) Characterization of Pulley Ridge 

Coral and Fish Fauna. Technical Report to 

the Gulf of Mexico Fisheries Management 

Council, Tampa, Florida. 72 pp.

Hamilton P (1990) Deep currents in the 

Gulf of Mexico. Journal of Physical 

Oceanography 20:1087-1104

Hamilton P, Lugo-Fernandez A (2001) 

Observations of high speed deep 

currents in the northern Gulf of Mexico. 

Geophysical Research Letters 28:2867-

2870.

Holmes CW (1981) Late Neogene and Quaternary 

geology of the southwestern Florida

shelf and slope. U.S. Geological Survey, 

Open File Report 81-1029, p27

Inoue M, Welsh SE, Rouse L, Weeks E (2002) 

Observation of deep water manifestation 

of Loop Current and Loop Current rings 

in the eastern Gulf of Mexico. EOS, 

Transactions, American Geophysical 

Union 52:321

Jordan GF, Stewart HB (1961) Submarine 

topography of the western Straits of Florida. 

Geological Society of America Bulletin 

72:1051-1058
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

GULF OF MEXICO

298

Jordan GF, Malloy RM, Kofoed JW (1964) 

Bathymetry and geology of the Pourtales 

Terrace. Florida. Marine Geology 1:259-

287.

Koenig CC, Coleman FC, Scanlon K (2000a) 

Reef fish resources and gas pipelines 

on the west Florida shelf: potential 

conflicts In: McKay M, Nides J, Vigil D 

(eds) Proceedings: Gulf of Mexico fish 

and fisheries: bringing together new and 

recent research, October 2000. U.S. 

Dept. of the Interior, Minerals Management 

Service, Gulf of Mexico OCS Region, 

New Orleans, La. OCS Study MMS 2002-

004.109-119

Koenig CC, Coleman FC, Grimes CB, Fitzhugh 

GR, Scanlon KM, Gledhill CT, Grace 

M (2000b) Protection of fish spawning 

habitat for the conservation of warmtemperate reef-fish fisheries of shelf-edge 

reefs in Florida. Bulletin of Marine Science 

66:593-616

Land L, Paull C (2000) Submarine karst belt rimming 

the continental slope in the Straits of Florida. 

Geo-Marine Letters 20:123-132

Leipper DF, Cochrane JD, Hewitt JF (1972) A 

detached eddy and subsequent changes. 

In: Capurro LRA, Reid JL (eds) Contributions 

on the physical oceanography of the Gulf 

of Mexico. Texas A&M Oceanographic 

Studies Vol. 2, Gulf Publishing Co., 

Houston, Texas, pp107-117

Lindberg WD, Lockhart FD (1993) Depth stratified 

population structure of geryonid crabs 

in the eastern Gulf of Mexico. Journal of 

Crustacean Research 13:713-722

MacDonald IR, Boland GS, Baker JS, Brooks 

JM, Kennicut II MC, Bidigare RR (1989) 

Gulf of Mexico hydrocarbon seep 

communities II Spatial distribution of 

seep organisms and hydrocarbons at 

Bush Hill. Marine Biology 101:125-128

Messing CG, Neumann AC, Lang JC (1990) 

Biozonation of deep-water Lithoherms 

and associated hardgrounds in the 

northeastern Straits of Florida. Palaois 

5:15-33

Meyer DL, Messing CG, Macurda Jr DB (1978) 

Zoogeography of tropical western Atlantic 

Crinoidea. Bulletin of Marine Science 

28:412-441

Minter LL, Keller G, Pyle T (1975) Morphology 

and sedimentary processes in and around 

Tortugas and Agassiz Sea Valleys, 

southern Straits of Florida. Marine 

Geology 18:47-69

Moore DR, Bullis Jr HR (1960) A deep-water 

coral reef in the Gulf of Mexico. Bulletin of 

Marine Science 10:125-128

Mortensen PB, Rapp HT (1998) Oxygen and 

carbon isotope ratios related to growth line 

patterns in skeletons of Lophelia pertusa 

(L) (Anthozoa, Scleractinia): implications 

for determination of linear extension rates. 

Sarsia 83:433-446.

National Marine Fisheries Service (1996) The 

Magnuson-Stevens Fishery Conservation 

and Management Act. United States 

Department of Commerce, National 

Oceanic and Atmospheric Administration, 

NMFS, Technical Memorandum NMFS­F/

SPO-23.

Newton CR, Mullins HT, Gardulski AF (1987) 

Coral mounds on the West Florida Slope: 

unanswered questions regarding the 

development of deepwater banks. 

Palaios 2:359-367

Nipper M, Sánchez Chávez JA, Tunnell Jr 

JW (eds) (2006) GulfBase: Resource 

Database for Gulf of Mexico Research. 

World Wide Web electronic publication. 

http://www.gulfbase.org

Nowlin WD (1971) Water masses and general 

circulation of the Gulf of Mexico. 

Oceanology 5(2):28-33
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

GULF OF MEXICO

299

Ostarello GL (1973) Natural history of the 

hydrocoral Allopora californica (Verrill 

1866). Biological Bulletin 145:548-564.

Parker NR, Mladenov PV, Grange KR (1997) 

Reproductive biology of the antipatharian 

black coral Antipathes fordensis in Doubtful 

Sound, Fjordland, New Zealand. Marine 

Biology 130:11-22

Parker Jr RO, Mays RW (1998) Southeastern 

U.S. deepwater reef fish assemblages, 

habitat characteristics, catches, and life 

history summaries. U.S. Dept. Commer. 

NOAA Tech. Rep. NMFS 138, p41

Paull CK, Neumann AC (1987) Continental 

margin brine seeps: their geological 

consequences. Geology 15:545-548

Paull CK, Chanton J, Martens C, Fullagar P, 

Neumann AC, Coston J (1991) Seawater 

circulation through the flank of the Florida 

Platform: evidence and implications. 

Marine Geology 102:265-279

Paull CK, Freeman-Lynde R, Bralower T, Gardemal 

J, Neumann AC, D’Argenio B, Marsella E 

(1990) Geology of the strata exposed on 

the Florida Escarpment. Marine Geology 

91:177-194

Pequegnat WE (1972) A deep bottom current 

on the Mississippi Cone. In: Capurro LRA, 

Reid JL (eds) Contributions on the physical 

oceanography of the Gulf of Mexico. 

Texas A&M Oceanographic Studies Vol. 

2, Gulf Publishing Co., Houston, Texas, 

pp107-117

Phillips NW, Gettleson DA, Spring KD (1990) 

Benthic biological studies of the southwest 

Florida shelf. American Zoologist 30:65-

75

Reed JK (2004) General description of deepwater coral reefs of Florida, Georgia and 

South Carolina: A summary of current 

knowledge of the distribution, habitat and 

associated fauna. A Report to the South 

Atlantic Fishery Management Council, 

NOAA, NMFS, p71

Reed JK, Wright A (2004) Submersible and 

scuba collections on deepwater reefs off 

the east coast of Florida, including the 

Northern and Southern Straits of Florida 

and Florida Marine Keys National Sanctuary 

for biomedical and biodiversity research of 

the benthic communities with emphasis 

on the Porifera and Gorgonacea, May 

20-June 2, 2004. Final cruise report. 

Conducted by the Center of Excellence, 

HBOI and FAU, p54. 

Reed JK, Wright A, Pomponi S (2004) Medicines 

from the deep sea: exploration of the 

northeastern Gulf of Mexico. Proceedings 

of the American Academy of Underwater 

Sciences 23th annual scientific diving 

symposium, March 11-13, 2004, Long 

Beach, California. pp58-70

Reed JK, Pomponi S, Wright A, Weaver D, Paull.

C (2005a) Deep-water sinkholes and 

bioherms of south Florida and Pourtales 

Terrace – habitat and fauna. Bulletin of 

Marine Science 77:267-296

Reed, J.K., D. Weaver, S.A. Pomponi. (2006a). 

Habitat and fauna of deep-water Lophelia 

pertusa coral reefs off the Southeastern 

USA: Blake Plateau, Straits of Florida, and 

Gulf of Mexico. Bulletin of Marine Science 

78:343-375.

Rezak R, Bright TJ, McGrail DW (1985) Reefs 

and Banks of the Northwestern Gulf of 

Mexico: Their Geological, Biological, 

and Physical Dynamics. John Wiley and 

Sons, New York 

Rogers AD (1999) The biology of Lophelia pertusa 

(Linnaeus 1758) and other deep-water 

reef-forming corals and impacts from 

human activities. International Review of 

Hydrobiology 84:315-406

Rowe GT, Kennicutt II MC (2001) Deepwater 

Program: Northern Gulf of Mexico 

Continental Slope Habitat and Benthic 

Ecology. Year I Interim Report. U.S. Dept. 

of the Interior, Minerals Management 

Service, Gulf of Mexico OCS Region, New 

Orleans, La. OCS Study MMS 2001-091. 

p674
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

GULF OF MEXICO

300

Sanders HL (1979) Evolutionary ecology and 

life history patterns in the deep sea. 

Sarsia. 64:1-7

Schmahl GP, Hickerson EL (2006) McGrail bank, 

a deep tropical coral reef community in 

the northwestern Gulf of Mexico. Proc 

10th International Coral Reef Symposium 

1:1124-1130

Schroeder WW (2002) Observations of Lophelia 

pertusa and the surficial geology at a 

deepwater site in the northeastern Gulf 

of Mexico. Hydrobiologica 471:29-33

Schroeder WW, Gittings SR, Dardeau MR, 

Fleischer P, Sager WW, Schultz AW, 

Rezak R (1989) Topographic features of 

the L’MAFLA continental shelf, Northern 

Gulf of Mexico. Proceedings Oceans ’89: 

The Global Ocean 1:54-59

Schroeder WW Brooke SD Olson JB Phaneuf 

B McDonough III JJ, Etnoyer, P (2005) 

Occurrence of deep-water Lophelia 

pertusa and Madrepora oculata in the Gulf 

of Mexico. Pages 297-307 in Freiwald 

A, Roberts JM (eds.), Cold-water corals 

and ecosystems. Springer-Verlag Berlin 

Heidelberg

Snyder D. (2000) Chapter 10: Fishes and Fisheries. 

In: Continental Shelf Associates (eds) 

Deepwater Gulf of Mexico environmental 

and socioeconomic data search and 

literature synthesis Volume 1: Narrative. 

Report US Dept. of the Interior, Minerals 

Management Service, Gulf of Mexico 

OCS Region, New Orleans LA. OCS study 

MMS 2000-049. pp255-279

Squires DF (1964) Fossil coral Thickets in 

Wairapa, New Zealand. Journal of 

Paleontology 38:904-915

Sulak KJ, Brooks RA, Luke KE, Norem AD, 

Randall M, Quaid AJ, Yeargin GE, Miller 

JM, Harden WM, Caruso JH, Ross SW. 

Demersal fishes associated with Lophelia 

pertusa coral and hard-substrate biotopes 

on the continental slope, northern Gulf of 

Mexico. Bulletin of Marine Science (in 

press)

Thayer CW (1989) Brachiopods and 

sclerosponges: dropouts from the 

evolutionary race. Geol Soc Amer, 1989 

Annual Meeting. pp252-253

Uchupi E (1968) Tortugas Terrace, a slip 

surface? U.S. Geological Professional 

Papers 600-D:D231-D234.

US Department of Energy (2006) An interagency 

roadmap for methane hydrate research 

and development. Prepared by the 

Technical Coordination Team, National 

Methane Hydrate R & D Program for 

the U.S. Department of Energy, Office of 

Fossil Energy. www.fossil.energy.gov

U.S. Department of the Interior, Minerals 

Management Service (2000) Gulf of 

Mexico deepwater operations and 

activities, environmental assessment. 

U.S. Department of the Interior, Minerals 

Management Service, Gulf of Mexico 

OCS Region, New Orleans, LA. OCS 

EIS/EA MMS 2000-001.

U.S. Department of the Interior, Minerals 

Management Service (2004) ‘Gulf 

of Mexico Deep Water Oil and Gas 

Production Remains Strong in 2003’ 

Press release #3103
STATE OF DEEP CORAL ECOSYSTEMS IN THE GULF OF MEXICO REGION

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Taxon Species Distribution Depth (m)* Reference

Phylum CnidariaClass AnthozoaSubclass HexacoralliaOrder ScleractiniaFamily Caryophyllidae Anomocora fecunda Pourtales 1871 SW Florida, N, NE GOM 73-779 Cairns 1978, 1987

Asterosmilia prolifera Pourtales 1871 N. GOM 32-311 Cairns 1978Caryophyllia ambrosia Alcock 1898 SW Florida, N, NE GOM 183-2360 Cairns 1978Caryophyllia berteriana Duchassaing 1850 SW Florida, N.GOM 100-850 Cairns 1978, 1987Caryophyllia cornuformis Pourtales 1870 SW Florida 137-931 Cairns 1978Caryophyllia parvula Cairns 1979 N GOM 68-136 Cairns 1979Coenosmilia arbuscula Pourtales 1871 SW Florida 109-807 Cairns 1978Dasmosmilia variegata Pourtales 1871 SW Florida 185-600 Cairns 1978Dasmosmilia lymani Pourtales 1871 N. GOM 33-366 Cairns 1978Deltocyathus italicus Michelotti 1838 SW Florida, N.GOM 403-2634 Cairns 1978, 1987Deltocyathus calcar Pourtales 1874 SW Florida, N.GOM 101-675 Cairns 1978Deltocyathus eccentricus Cairns 1979 SW Florida, N.GOM 183-910 Cairns 1978Desmophyllum dianthus Esper 1794 NE SE GOM 900-1210 NMNHLophelia pertusa (Linnaeus, 1758) SW Florida, N. GOM, Wrecks 60-2170 Cairns 1978Oxysmillia rotundifolia Milne Edwards and Haime 1849 N. GOM 46-640 Cairns 1978Paracyathus pulchellus Philippi 1842 SW Florida, N.GOM, Wrecks 25-838 Cairns 1978, TAMUPepnocyathus stimpsonii Pourtales 1871 SW Florida 110-293 Cairns 1978Pourtalosmilia conferta Cairns, 1978 Wrecks 87-554 Schroeder pers. comm.Solenosmilia variabilis Duncan, 1873 Florida Straits 220-3383 Cairns 1978, Reed et al. 2005bStephanocyathus diadema Moseley 1876 N. GOM 795-2133 Cairns 1978Stephanocyathus paliferus Cairns 1977 SW Florida 229-1158 Cairns 1978

Appendix 7.1 Species list of deep coral species found in the Gulf of Mexico. Outlined/highlighted text indicates major structure forming species. Depth ranges are global depth ranges extracted from the literature. National Musuem of Natural History (NMNH), Texas A&M University (TAMU)
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Taxon Species Distribution Depth (m) Reference

Stephanocyathus coronatus Pourtales 1867 N. GOM 543-1280 Cairns 1978Thalamophyllia riisei Duchassaing and Michelotti 1860 N. GOM 18-1317 Cairns 1979, 1987Theocyathus laevigatus Pourtales 1871 SW Florida 183-576 Cairns 1978Trochocyathus flos Pourtales 1878 N. GOM 22-560 Cairns 1978Trochocyathus rawsonii Pourtales 1874 SW Florida 131-622 Cairns 1978

Family Dendrophylliidae Balanophyllia palifera Pourtales 1878 N. GOM 53-708 Cairns 1978, 1987

Balanophyllia floridana NE GOM 92m Wicksten pers. com.Bathypsammia tintinnabulum Pourtales 1868 SW Florida 210-1079 Cairns 1978Bathypsammia? sp. Marenzeller, 1907 SW Florida 418-640 Reed et al. 2005bDendrophyllia cornucopia Pourtales 1871 SW Florida 132-960 Cairns 1978Dendrophyllia alternata Pourtales 1880 N. GOM 276-1200 Cairns 1978Enallopsammia profunda Pourtales 1867 SW Florida 146-1748 Cairns 1

978

Rhizopsammia manuelensis Chevalier 1966 N. GOM 55-366 Cairns 1978

Family Fungiacyathidae Fungiacyathus crispus Pourtales 1871 SW Florida, N.GOM 183-1010 Cairns 1978Family Flabellidae Flabellum moseleyi Pourtales 1880 SW Florida, N. GOM 216-1097 Cairns 1978

Flabellum fragile Cairns 1977 SW Florida 80-366 Cairns 1978Javania cailleti Duchassaing and Michelotti 1864 SW Florida, N. GOM 86-2165 Cairns 1978Rhizotrochus fragilis Pourtales 1868 SW Florida 90-796 Cairns 1978

Family Guyniidae Guynia annulata Duncan 1872 SW Florida, N. GOM 3-653 Cairns 1978

Schizocyathus fissilis Pourtales 1874 SW Florida, N. GOM 88-640 Cairns 1978Stenocyathus vermformis Pourtales 1868 SW Florida 128-1229 Cairns 1978

Family Oculinidae Madrepora oculata Pourtales 1758 N. GOM 80-1500 Cairns 1

978

Madrepora carolina Pourtales 1871 SW Florida, N.GOM 53-1003 Cairns 1

978

Oculina varicosa Lesueur, 1821 Wrecks 87 Schroeder pers. com.

Family Pocilloporidae Madracis brueggemanni Ridley N GOM 55-127 NMNH

Madracis myriaster Milne Edwards and Haime 1849 SW Florida, N.GOM, Wrecks 37-875 Cairns 1978Madracis formosa Wells 1973 SW Florida 46-104 NMNH
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Taxon Species Distribution Depth (m) Reference

Madracis pharensis SW Florida 90 Wicksten pers. com.

Order Antipatharia

Acanthopathes thyoides N. GOM 124 FGBNMS/OpreskoAntipathes atlantica N. GOM 77 FGBNMS/OpreskoAntipathes sp. cf. gracilis N. GOM 70 FGBNMS/OpreskoAntipatharia, unid. Florida Straits, N. GOM 283-767 Reed et al. 2005bAntipatharia, unid. Florida Straits, SW Florida 328-515 Reed et al. 2005bAntipathes columnaris Duchassaing 1870 SW Florida 73-567 Cairns 1993Antipathes furcata Gray, 1857 Wrecks, N.GOM 87-106 FGBNMS/OpreskoAntipathes rigida Pourtales, 1880 Florida Straits, S, N GOM 64-640 Cairns 1993, Reed et al. 2005bAntipathes pedata Gray 1857 SW Florida, N. GOM 60-308 Cairns 1993Antipathes tanacetum Pourtales 1880 Florida Straits, N. GOM 46-915 Cairns 1993Antipathes salix Pourtales 1880 SW Florida, N. GOM 107-263 Cairns 1993Aphanipathes abietina Pourtales 1874 SW Florida, N. GOM 31-310 Cairns 1993Aphanipathes pedata N. GOM 131 FGBNMS/OpreskoBathypathes patula Brook 1889 SW Florida 100-5000 Cairns 1993Bathypathes alternata Brook, 1889 SW Florida 466-716 Reed et al. 2005bElatopathes abietina N. GOM 106 FGBNMS/OpreskoLeiopathes glaberrima Esper., 1786 N. GOM 176-549 Cairns 199

3

Phanopathes expansa N. GOM 121 FGBNMS/OpreskoPlumapathes pennacea N. GOM 61 FGBNMS/OpreskoStichopathes sp. cf. S. pourtalesi Brook, 1889 Wrecks 87 Schroeder pers. com.Stichopathes lutkeni N. GOM 64 FGBNMS/OpreskoTanacetipathes barbadensis N. GOM 89 FGBNMS/OpreskoTanacetipathes tanacetum N. GOM 107 FGBNMS/Opresko

Subclass OctocoralliaOrder AlcyonaceaFamily Alcyoniidae Anthomastus (Bathyalcyon) robustus delta Bayer 1993 N GOM 274 NMNH

Anthomastus sp. Florida Straits NMNH
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Taxon Species Distribution Depth (m) Reference

Family Clavulariidae Telesto flavula Deichmann 1936 N GOM

Telesto fruticulosa Dana N GOM

Family Nidaliidae Nidalia occidentalis Gray 1835 N GOM 76-214 NMNH

Siphonogorgia agassizii Deichmann 1936 N GOM 58-190 NMNHChironepthya (Siphonogorgia) caribaea Deichmann 1936 NW GOM 75-135 NMNH/FGBNMS

Family Plexauridae Bebryce cinerea N GOM 107 FGBNMS/Williams

Bebryce sp. cf. cimeria N GOM 91 FGBNMS/EtnoyerBebryce parastellata Deichmann 1936 N GOM 46-586 NMNHCaliacis nutans Duchassaing & Michelotti, 1864 SW Florida, N GOM 101-238 NMNHEchinomuricea atlantica Johnson, 1862 SW Florida 101-183 NMNHHypnogorgia pendula Duchassaing & Michelotti, 1864 N GOM 60-183 NMNHMuricea pendula Verrill, 1864 Wrecks, N GOM 53-87

Schroeder pers. com., FGBNMS/Cairns

Muriceides sp. cf. furta N. GOM 91 FGBNMS? EtnoyerMuriceides sp. Florida Straits 191 Reed et al. 2005bParamuricea multispina Deichmann, 1936 Florida Straits 176 NMNHParamuricea placomus Linnaeus, 1924 Florida Straits 462-470 Reed et al. 2005bParamuricea multispina Deichmann, 1936 Florida Straits 189-715 Reed et al. 2005bPlacogorgia tenuis Verrill, 1883 Florida Straits, N. GOM 86-557 NMNH, Reed et al. 2005b, EtnoyerPlacogorgia mirabilis Deichmann, 1936 Florida Straits, N. GOM 172-212 Reed et al. 2005bPlacogorgia sp. (red unidentified) Placogorgia rudis Florida Straits, Wrecks 143-238 NMNH, Schroeder pers. com.Plumarella pourtalesi Verrill, 1883 Florida Straits 171-753 Reed et al. 2005bScleractis guadaloupensis Duchassaing & Michelotti, 1860 N. GOM 58-190Swiftia casta Verrill, 1883 Florida Straits 53-942 NMNH, Reed et al. 2005b
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Taxon Species Distribution Depth (m) Reference

Swiftia new sp.? Florida Straits 497 Reed et al. 2005bThesea sp. cf. T. grandiflora Deichmann, 1936 SW Florida, N GOM, Wrecks 143 NMNH, Schroeder pers. com.Thesea sp. cf. T. rubra Deichmann, 1936 N GOM, Wrecks 61-143 NMNH, Schroeder pers. com.Thesea sp. Wrecks 143 Schroeder pers. com.Trachymuricea hirta Pourtales, 1867 Florida Straits 462 Reed et al. 2005bThesea parviflora Deichmann, 1936 Florida Straits 183 Reed et al. 2005bVillogorgia nr. nigrescens Duchassaing & Michelotti, 1860 Florida Straits 215 Reed et al. 2005b

Order GorgonaceaFamily Anthothelidae Diodogorgia nodulifera Hargit in Rogers and Hargitt 1901 N. GOM 86 FGBNMS/EtnoyerFamily Chrysogorgiidae Chrysogorgia elegans SW Florida, N. GOM 162-732 NMNH

Chrysogorgia spiculosa Florida Straits, N. GOM 1256-2360 NMNH

Family Ellisellidae Ellisella atlantica (Toeplitz) N GOM 56-100 NMNH

Ellisella barbadensis Duchassaing & Michelotti Florida straits, NW, N GOM 64-390 NMNHEllisella funiculina Duchassaing & Michelotti NE GOM 55-92 NMNHEllisella elongata Pallas NE GOM 55-91 NMNHNicella deichmani N. GOM 107 FGBNMS/CairnsNicella guadalupensis Duchassaing & Michelotti 1846 N GOM 68-159 NMNHNicella goreaiu N. GOM 86 FGBNMS/EtnoyerRiisea paniculata Duchassaing & Michelotti 1860 N GOM 188 NMNH

Family Gorgoniidae Leptogorgia stheno Bayer N GOM 69 NMNHFamily Isididae Acanella arbuscula SW Florida, NW GOM 940m NMNH

Acanella sp. GOM 183-1216 m Wicksten pers. com.Keratoisis flexibilis Pourtales 1868

N GOM, SW Florida, Florida Straits 378-816 NMNH
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Taxon Species Distribution Depth (m) Reference

Family Keroeididae Thelogorgia stellata Bayer 1992 N. GOM 89 FGBNMS/CairnsFamily Primnoidae Callogorgia americana delta Cairns and Bayer, 2002 N GOM 366-570 Cairns and Bayer 2002

Callogorgia americana americana Cairns and Bayer, 2002 Florida Straits 183-732 Cairns and Bayer 2002Callogorgia gracilis Milne Edwards and Haime, 1857 N. GOM 110 FGBNMS/Cairns

Order Pennatulacea

Umbellula guntheri Kolliker, 1880 N GOM 2683 NMNHUmbellula lindahlii Kolliker, 1874 N GOM 2683 NMNHFuniculina quadrangularis Pallas N GOM 768 NMNH

Class HydrozoaOrder AnthoathecataeFamily Stylasteridae Stylasteridae, unid. sp. Florida Straits 173-742 Reed et al. 2005b

Stylaster erubescens Pourtales, 1868 Florida Straits 175-186 Reed et al. 2005bStylaster miniatus Pourtales, 1868 Florida Straits 175-220 Reed et al. 2005b, NMNHStylaster filogranus Pourtales, 1871 Florida Straits 175-366 Reed et al. 2005b, NMNHStylaster unid. sp.1 Florida Straits 557 Reed et al. 2005bStylaster laevigatus Cairns, 1986 Florida Straits 366 NMNHStylaster aurantiacus Cairns, 1986 SW Florida 159-166 NMNHStylaster gemmascens Florida Straits 183 NMNHDistichopora foliacea Pourtales, 1868 Florida Straits 175 Reed et al. 2005bCrypthelia glossopoma Cairns, 1986 Florida Straits 205-210 NMNHPliobothrus echinatus Cairns, 1986 Florida Straits 175 NMNH, Reed et al. 2005b* Note: Depth ranges are not just GOM samples, but are global depth ranges extracted from the literature
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STATE OF DEEP CORAL ECOSYSTEMS

IN THE CARIBBEAN REGION:

PUERTO RICO AND THE U.S. VIRGIN ISLANDS

Steven J. Lutz1,2 and Robert N. Ginsburg1

1

Ocean Research and Education Foundation

1300 Galiano St., Coral Gables, FL 33134

2

Corresponding author addres: Marine Conservation Biology Institute, Public Policy Office

600 Pennsylvania Ave SE, Suite 210,

Washington DC 20003 

I. INTRODUCTION 

This chapter summarizes published and 

unpublished information on deep corals in the 

wider Caribbean region (Caribbean Sea and 

Bahamian Archipelago) including those in the 

exclusive economic zone (EEZ) of United States 

Possessions. It summarizes the occurrences of 

the corals, associated species and their sea floor 

accumulations; it provides basic information on 

the geography, geology and oceanography of the 

region and how they interact to produce sea floor 

elevations; and it discusses stressors, including 

potential fisheries interactions. 

For the purposes of this chapter deep corals are 

defined as those in waters below 100 m. 

The territorial jurisdiction of the U.S. in the 

Caribbean is limited to the areas around Puerto 

Rico, the U.S. Virgin Islands and the uninhabited 

island of Navassa between Cuba and Hispaniola, 

illustrated in Figure 8.1. The EEZ of Puerto Rico 

and the U.S. Virgin Islands extends from the 

edge of the territorial waters to 200 nautical miles 

to the north and is bounded to the south by the 

EEZ of Venezuela, to the east by the British Virgin 

Islands and to the west by Dominican Republic. 

The EEZ of Navassa Island is bounded to the 

north by Cuba, to the west by Jamaica and to the 

east by Haiti.

As information on deep-sea corals in the U.S. 

Caribbean is limited, a consideration of the wider 

Caribbean enables a broader understanding of 

reported and potential deep water corals in the 

U.S. Caribbean.

Regional Deep-sea Research

Louis F. de Pourtalès first discovered deep-sea 

life in the region, including deep-water corals, 

in the 1860’s. Pourtalès, a scientist aboard the 

Coast Survey steamer Bibb, found evidence of life 

off Cuba at 945 m (3,100 ft). He studied regional 

deep-sea corals and published comprehensive 

works (Pourtalès 1867, 1871). Pourtalès was 

also a pioneer of deep-sea dredging, a method 

that was used by later research cruises. These 

cruises were not focused on deep-sea corals and 

recovered specimens were incidental findings. 

Since the discoveries made by Pourtalès and 

the Bibb many research cruises have contributed 

to the understanding of deep-sea corals. Early 

cruises included voyages by the U.S. Coast 

Survey vessel Blake (late 1870’s), U.S. Fish 

Commission Albatross (1880’s) and the H.M.S. 

Challenger (1872-1876). Later research cruises 

included voyages by the R/V Oregon and R/V 

Oregon II (1950’s to 1970’s), R/V Gerda (1960’s), 

R/V Pillsbury (1960’s and 70’s), the R/V Eastward

(1970’s), the R/V Akademik Vernadski (early 

1970’s); and R/V Columbus Iselin (1970’s and 

80’s).

Beginning in the 1970’s, dredging was 

supplemented by the use of research 

submersibles. These direct observations and 

photographs of deep-sea coral communities 

provided a major advance in understanding the 

occurrences, morphology and communities of 

deep-sea corals in the Straits of Florida and off 

the northeast slope of the Little Bahama Bank 

(Figure 8.1) (Neumann and Ball 1970; Neumann 

et al. 1977; Messing 1990; Reed et al. 2005). 

Other submersible research of the region includes 

the following: to 300 m (the “sub reef habitat”) 

at Discovery Bay Jamaica (Lang 1974; Hartman 

1973; Lang et al. 1975); to 300 m in Belize 

(Ginsburg and James 1979); research by Harbor 

Branch Oceanographic Institute (HBOI) to 1,000 

m for sites in the Greater and Lesser Antilles 

and Netherlands Antilles (Reed pers. comm.). 
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Figure 8.1. Study area showing geography and United States territories. The approximate boundaries of U.S. Caribbean EEZs are identified by the blue dashed line: A) Puerto Rico and the U.S. Virgin Islands; B) Navassa Island. 
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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Figure 8.2. Regional currents, channels, and passages (adapted from Gyory et al. 2005a and b).

Submersible research for the U.S. Caribbean 

includes dives by HBOI and NOAA (Nelson and 

Appeldoorn 1985).

In 1979 Stephen Cairns published a seminal 

review of deep-water stony corals occurring in the 

Caribbean Sea and adjacent waters, examining 

eighty-eight species of deep-water corals (both 

solitary and colonial species) (Cairns 1979). 

Recent regional deep-sea, coral related, 

investigations include the following: research on 

the distribution of deep water coral mounds in 

the Straits of Florida by the use of high-resolution 

multibeam sounding (Correa et al. 2006; McNeill 

2006); continued private pharmaceutical research 

throughout the region and especially in the 

Straits of Florida interests (Reed pers. comm.); 

research on azooxanthellate coral communities 

in Colombia’s Caribbean waters, off the San 

Bernardo-Rosario and Santa Marta Islands 

(Reyes et al. 2005; Santodomingo et al. 2006); 

and an ongoing effort to characterize seafloor 

habitats down to 1,000 meters within the U.S. 

Virgin Islands and Puerto (NOAA 2007).

II. GEOLOGICAL SETTING

The Caribbean Sea basin, the islands within 

it, and the Bahamas Archipelago, consisting of 

shallow banks and inter-bank depressions, make 

up this region (Figure 8.1). While most of the 

area is centered on the Caribbean plate several 

continental plates meet in this region, which are 

hotspots for seismic activity. Interactions between 

the Caribbean plate, North American plate and 

the South American plate produced the major 

topographic features and allow the Caribbean 

basin to be divided into four smaller basins; the 

Yucatan, Colombian, Venezuelan and Granada 

basins (Figure 8.1). 

Both tectonic activity and biological activity 

produce features significant to deep water corals 

in this region. Tectonic activity is responsible for 

deep water trenches, ridges, basins and interisland passages and channels (topographic 

features are detailed in section III). Products of 

biological activity include deep coral mounds and 

lithoherms (discussed in section IV). 
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III. OCEANOGRAPHIC SETTING

The Caribbean Sea is approximately 2,640,000 

km2 or 1,425,485 square nautical miles. It is a 

semi-enclosed basin, bounded on the west by 

Central America, on the south by Central and 

South America, on the east by the Lesser Antilles 

and on the north by the Greater Antilles. With 

a few exceptions, the Caribbean Sea generally 

exceeds 1,830 m (6,000 ft) in depth with many 

sections exceeding 3,660 m (12,000 ft) in depth. 

Its greatest known depth is 7,535 m (24,721 ft) 

below sea level in the Cayman Trench located 

between Cuba and Jamaica. The Puerto Rican 

Trench, the deepest area within the study region 

and deepest part of the Atlantic Ocean, lies north 

of Puerto Rico just outside the Caribbean, with a 

depth of 9,200 m (30,183 ft). The only notable 

extensive shallow areas are the Bahamas Banks 

and the Nicaragua Rise. Several gaps between 

islands on the north and east are major channels 

and passes for ocean currents connecting the 

Caribbean Sea with the Atlantic Ocean and Gulf 

of Mexico. Chief among these are the Windward 

Passage between Cuba and Hispaniola; the 

Mona Passage between Puerto Rico and 

Hispaniola; the Anegada Passage south of the 

Virgin Islands; smaller channels between islands 

in the Windward Island chain; and the Grenada 

Passage between the Lesser Antilles and the 

South American continent (Figure 8.2).

As a result of the exchanges with the open Atlantic, 

there is little seasonal variation in surface water 

temperatures, with a general range from 25.5o C 

in the winter to 28 o C in the summer. Below the 

surface, water structure is highly stratified in the 

upper 1200 meters; weakly stratified between 

1,200 and 2,000 meters and uniform below 2000 

meters. This stratification is directly related to the 

shallow depth of passages between the Lesser 

Antilles. They act as sills and impede deep-water 

flow into the Caribbean (Gordon 1967; Gyory et 

al. 2005a). 

Most water flows into the Caribbean Sea from 

the Atlantic through the Grenada, St. Vincent and 

St. Lucia Passages in the southeast (Figure 8.2) 

(Johns et al. 2002; Gyory et al. 2005a). From 

these passages, water flows clockwise across the 

Caribbean Sea basin as the Caribbean Current, 

the main surface circulation in the Caribbean 

Sea. It reaches high surface velocity (70 cm/

sec) along the Netherlands Antilles and adjacent 

coastline of South America (Fratantoni 2001). 

The current then flows to the northwest, over the 

Colombian basin, towards the Nicaragua Rise 

and a trough southwest of Jamaica, with a branch 

forming an counterclockwise Panama-Colombia 

Gyre (where the current meets Central America). 

Strong flow (up to 60 cm/sec) has been recorded 

along the Panamanian and Colombian coasts 

(Gyory et al. 2005a).

Where currents are diverted by or constricted 

between landmasses, current velocities increase. 

The larger and deeper inter island spaces produce 

major seaways; the more numerous, smaller 

and shallower ones produce similar but smaller 

increases in velocity. Current velocities will 

also increase when water flow meets elevations 

of the sea floor, typically platforms or banks of 

calcareous deposits. For example, the passages 

between the large and long-lived platforms of the 

Bahamian Archipelago and the Florida Peninsula 

are sites of increased high flow. 

At the Nicaragua Rise channel-like constrictions 

between shallow banks accelerate current. The 

flow and water is accelerated through the trough 

southwest of Jamaica and then turns west as it 

crosses the Cayman Ridge and enters Yucatan 

Channel as the Yucatan Current (Gyory et al. 

2005d). Strong currents (170 cm/sec) have been 

reported in the Yucatan Channel (Pillsbury 1890; 

Gyory et al. 2005d). Water flow exits the Yucatan 

Channel into the Gulf of Mexico and doubles back 

as the Loop Current (Gyory et al. 2005c) before 

entering the Straits of Florida between Western 

Great Bahama Bank and the Florida Reef Tract 

where it is joined by waters passing through the 

Old Bahama Channel to form the Florida Current 

(Gyory et al. 2005b). Gyory et al. (2005b) reports 

strong surface flow in the Florida Current; “at 

times there is a 2 m/sec flow within a few miles 

of the (Florida) coast.” When this Florida Current 

exits the Straits it is joined by the Antilles Current 

and turns eastward and becomes the Gulf Stream 

sensu stricto. 

The Antilles Current transports tropical waters 

from the Atlantic northwestward, flowing north 

of the Greater Antilles and joins the Gulf Stream 

off Florida past the outer Bahamas (Figure 8.2) 

(Rowe et al. 2005). Its waters are concentrated 

into a strong northward jet centered at 400 meters 

deep about 80-100 km wide (Lee et al. 1996). It 

is a considerable source of warm water for the 
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Figure 8.3. Distribution of major and potential structure-forming stony corals (scleractinia) and deep sea bio-buildups (lithoherms, mounds, and banks). The letter S identifies the locations of trawl stations referred to in Appendix 8.2 (some locations overlap). References are listed in Appendix 8.1.
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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Figure 8.4. Reported distribution of potential habitat-forming corals (larger gorgonians, etc.). References include CITES 2007 and those listed in Appendix 8.1.
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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Gulf Stream system. However, it occurs more 

as an eddy field along the Bahamas Archipelago 

rather than as a constant flow (Gunn and Watt 

1982).

IV. STRUCTURE AND HABITAT-FORMING

DEEP CORALS

A wide variety of deep-sea corals occur within 

the wider Caribbean region. These include stony 

corals, gorgonians, soft corals, stylasterids, 

black corals, lithotelestid coral and sea pens. 

The knowledge about the distribution of these 

corals is generally considered poor, greater for 

some groups than others. Cairns (1979) notes, 

with regard to deep-sea stony corals, that the 

southern coasts of the Greater Antilles (Cuba and 

Hispaniola) and Costa Rica are not well sampled 

and that the Straits of Florida (including the 

northwest coast of Cuba) and the Lesser Antilles 

are well sampled. Many species of deep-sea 

corals occur within the U.S. Caribbean. Species 

with an Antillian distribution (occurring within the 

Greater and Lesser Antilles) may also occur in 

U.S. Caribbean waters. 

Deep-sea corals of interest to this chapter, are 

those termed “structure-forming” and “potential 

habitat-forming”. Structure-forming corals are 

associated with deep-sea coral habitat such as 

coral banks, bioherms and lithoherms. Both 

structure and non-structure forming corals occur 

within the study area (Figures 8.3, 8.4 and 

8.5). Bayer (1961) describes the importance of 

octocorals (gorgonians and soft corals) in the reef 

building process and framework of shallow reefs. 

The importance of octocorals in deep water reefs 

is not yet understood but may be great in terms of 

contribution to framework and baffling of marine 

sediment, larger octocorals are expected to 

contribute and baffle more than smaller species. 

Appendix 8.1 details the attributes of each coral 

species including distribution.

a. Stony corals (Class Anthozoa, Order 

 Scleractinia)

Ninety-nine species of azooxanthellate hard 

corals are reported from the study area in depths 

below 100 m. Most are small single polyp species, 

shaped like cups, which often occur solitarily (e.g. 

Figure 8.6). Solitary corals are not reported to form 

significant structures. Nine species of structureFigure 8.5. Distribution of deep water features and corals found in the U.S. Caribbean and adjacent area. 

References include EarthRef.org 2007 and those listed in Appendix 8.1.
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Table 8.1. Structure-forming deep water stony corals of the wider Caribbean region. The measurements given 

in brackets correspond to the approximate horizontal length for each specimen illustrated.

Species Description Illustration

Dendrophyllia 

alternata

D. alternata occurs throughout the North Atlantic. This coral 

forms bushy colonies up to 1 m in height and has been 

described as a reef builder. It occurs at depths of 279 - 900 

m in the tropical western Atlantic. Regionally, it has been 

recorded with M. oculata and M. myriaster (Cairns 1979; 

Cairns and Stanley 1982; Le Goff-Vitry 2003). [30 cm, 

specimen recovered at 290.8 m off Curacao]

Desmophyllum 

dianthus

D. dianthus (also known as D. cristagalli) forms densely 

branched colonies is reported to contribute to reef formation. It 

occurs globally to 2,460 m and in the tropical western Atlantic 

from 155 - 1,939 m. Regionally, it has been recorded with 

E. profunda, L. pertusa, M. oculata, and S. variabilis (Cairns 

1979; Cairns and Stanley 1982; Rogers 1999). [10.8 cm, 

specimen recovered at 597 m off the east coast of Florida]

Enallopsammia 

profunda

E. profunda occurs globally and is endemic to the western 

Atlantic. It forms massive bushy colonies, up to 1 m thick, 

and is regarded a major structure-forming species. It occurs 

from 403 - 1,748 m and is often associated with L. pertusa, 

M. oculata and S. variabilis (Cairns 1979; Cairns and Stanley 

1982; Rogers 1999; Reed 2002; Freiwald et al. 2004). [30.7 

cm, specimen recovered at 801 m in the Straits of Florida off 

Cuba]

Enallopsammia 

rostrata

E. rostrata occurs in the U.S. Caribbean and globally. This 

coral forms massive bushy colonies and is regarded a major 

structure-forming species. It occurs from 300 - 1,646 m in the 

tropical western Atlantic. It is associated with L. pertusa on 

deep-water reefs in the north Atlantic and has been recorded 

with S. variabilis, M. oculata, and E. profunda in the wider 

Caribbean (Cairns 1979; Rogers 1999; Reed 2002; Freiwald 

et al. 2004). [31.4 cm, specimen recovered at 1,097 m off 

Bermuda. Specimen includes L. pertusa and D. dianthus]

Lophelia 

pertusa

L. pertusa occurs in the U.S. Caribbean, globally, and is well 

known from the north Atlantic. This coral forms bushy colonies 

up to a meter in height and is regarded the most common 

reef-building deep/cold-water coal. It has been reported from 

waters as shallow as 25 m to as deep as 3,383 m. Regionally, 

it is most common between 500 - 800 m. L. pertusa is often 

found associated with E. profunda and has been recorded 

with D. dianthus, M. carolina, M. oculata, and S. variabilis in 

the wider Caribbean (Cairns 1979; Cairns and Stanley 1982; 

Rogers 1999; Dawson 2002; Reed 2002; Freiwald et al. 

2004). [24.3 cm, specimen recovered at 793 m in the Straits of 

Florida off Cuba]

building and branched azooxanthellate hard corals 

are reported from the study area: Dendrophyllia 

alternata, Desmophyllum dianthus (also known 

as Desmophyllum cristagalli), Enallopsammia 

profunda, Enallopsammia rostrata, Lophelia 

pertusa, Madracis myriaster, Madrepora oculata, 

Madrepora carolina and Solenosmilia variabilis

(Cairns pers. comm.). Table 8.1 and Figures 8.7 

- 8.10 and 8.15 describe and illustrate structurebuilding corals.

Six species of azooxanthellate hard corals 

are regarded as the major structure-forming 

species: E. profunda, E. rostrata, L. pertusa, M. 

carolina, M. oculata and S. variabilis. Two genera 

(Lophelia and Solenosmilia) constitute two of 
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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315

Species Description llustration

Madracis 

myriaster

M. myriaster occurs in the U.S. Caribbean and throughout 

the wider Caribbean. It forms bushy colonies and is reported 

as a main framework builder of deep-water coral banks in 

Colombian waters. Regionally, it has been recorded with M. 

carolina. It occurs from 37 - 1220 m in the tropical western 

Atlantic (Cairns 1979; Reyes et al. 2005; Santodomingo et al. 

2006). [30.2 cm, specimen recovered at 200 m off Jamaica]

Madrepora 

carolina

M. carolina occurs in the U.S. Caribbean and throughout the 

tropical western Atlantic. This coral forms bush-like colonies 

with a thick base and is associated with cold-water reefs 

as a structure-forming species. It occurs from 53 - 801 m . It occurs from 53 - 801 m 

(most commonly between 200 - 300 m). Regionally, it has 

been recorded with E. profunda, E. rostrata, and D. dianthus 

(Cairns 1979; Dawson 2002; Freiwald et al. 2004). [27.6 

cm, specimen recovered at 333 - 375 m in the northwest 

Providence Channel off Grand Bahama Island]

Madrepora 

oculata

M. oculata occurs in the U.S. Caribbean and globally. This 

coral forms large bushy colonies and is regarded a major 

structure-forming species. It occurs from 55 - 1,950 m. It 

is commonly associated with L. pertusa in the eastern and 

northeastern Atlantic and Mediterranean Sea. Regionally, it 

has been recorded to occur with E. profunda, E. rostrata, D 

dianthus, L. pertusa, and S. variabilis (Cairns 1979; Rogers 

1999; Dawson 2002; Reed 2002; Freiwald et al. 2004). [24.1 

cm, specimen recovered at 634 m in the Gulf of Mexico]

Solenosmilia 

variabilis

S. variabilis occurs globally. This coral forms bushy colonies 

and is regarded a major structure-forming species. It occurs 

from 220 - 2,165 m and is known from the northern slope 

of the Little Bahama Bank from 1,000 - 1,300 m, where it is 

associated with deep-water banks. It is often found associated 

with L. pertusa, Madrepora spp., and E. profunda, and has 

been recorded with E. rostrata and D. dianthus in the wider 

Caribbean (Cairns 1979; Mullins et al. 1981; Cairns and 

Stanley 1982; Rogers 1999; Freiwald et al. 2004). [20.7 cm, 

specimen recovered at 1,000 m in the North Atlantic Ocean]

Figure 8.6. Assorted solitary deep water stony corals: a) Stephanocyathus diadema, from The 

Bahamas at 1,779 m; b) S. laevifundus, from St. Vincent at 576-842 m; c) Caryophyllia ambrosia 

caribbeana, from Surinam at 1,318-1,343 m; d) Deltocyathus eccentricus, from Jamaica at 457-558 

m; e) Bathypsammia tintinnabulum, from the Straits of Florida at 689-699 m (B. tintinnabulum is not 

known from the region but is presented here for comparison).
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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316

Figure 8.9. Close up of the potential structureforming coral D. alternata, sampled by submersible, 

from the Netherlands Antilles, at 290.8 m.

reported from the U.S. Caribbean.

Another major structure-forming species, Oculina 

varicosa, occurs throughout the Caribbean. O. 

varicosa is a facultative zooxanthellate species 

(i.e. it occurs with zooxanthellate in shallow 

water and without in deeper water). O. varicosa

is reported to 152 m and forms bushy colonies 

several meters in height (commonly at 45 - 91 

m). It is described as a main framework builder 

of deep-water banks off Florida’s central eastern 

coast. However, as no deep water banks of 

O. varicosa have been reported from the wider 

Caribbean (only shallow water species) this 

species is not considered further.

Madracis myriaster, an azooxanthellate stony 

coral found throughout the wider Caribbean, 

is reported to form bushy shaped colonies and 

significantly contribute to habitat formation on 

the four major constituents of known world-wide 

deep-water coral banks at depths of 400 - 700 m 

(Cairns and Stanley 1982). Both M. oculata and 

M. carolina are regarded as minor constituents of 

coral banks in the Straits of Florida (Cairns 1979). 

Solenosmilia is the dominant structure-building 

coral found on deep coral banks north of the Little 

Bahama Bank (Reed 2002) (Table 8.2, Mullins 

Reefs), where Madrepora and Enallopsammia

also occur. Lophelia and Enallopsammia are 

common components of the lithoherms east of 

the Little Bahama Bank in the Straits of Florida 

(Reed 2002) (Table 8.2, Bahamas Lithoherms). 

Four major structure-forming species E. rostrata, 

L. pertusa, M. oculata and M. carolina have been 

Figure 8.8. Structure-forming coral (E. profunda) 

providing habitat for unidentified crustaceans and a 

crinoid. Photographed from submersible at 2,721 m 

in Straits of Florida. Photo credit: Reed 2006

Figure 8.10. Close up of the major structure-forming 

coral M. oculata, sampled by submersible from the 

Straits of Florida at 746.4 m. Photo credit: Reed 

2006.

Figure 8.7. Deep sea coral habitat (M. oculata and 

Paramuricea sp.) at 2,230 m in the Bahamas (Straits 

of Florida off Cat Cay). Photo credit: Reed 2006.
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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317

Location Description

Mullins Reefs

This site is characterized by a region of deep coral mounds (bioherms) on the northern 

deep shelf edge of the Little Bahama Bank at depths of 1000-1300 m. Mounds 5-40 m 

in height are reported. The bioherms are dominated by the major structure forming deep 

water coral Solenosmilia sp., Lophelia is reportedly absent. An area of approximately 1500 

km2 is described. Site profiled via. submersible, seismic reflections, and dredges (Mullins et 

al. 1981; Reed 2002).

Bahamas 

Lithoherms

This site is characterized by a region of lithoherms on the western shelf edge of the Little 

Bahama Bank in the Straits of Florida at depths of 300-500 m. Pinnacles 30-50 m in height 

are reported. The lithoherms are dominated by the major structure forming deep water 

coral L. pertusa. E. profunda and M. oculata are also present. D. alternata, A. fecunda, and 

the larger black coral Distichopathes filix have been reported from the region. An idealized 

lithotherm from this site is illustrated in Figure 8.23. Wreckfish have been reported in 

association with these lithoherms. An area of approximately 330 km2 is described. Site 

profiled via. submersible, seismic reflections, and dredges (Neumann and Ball 1970; 

Neumann et al. 1977; Messing et al. 1990; Reed 2002).

Mullins 

Mounds

Deep-water mounds (banks and lithoherms) and cemented hardgrounds are described off 

Bimini and entering the New Providence Channel. The mounds are described to continue 

northward and connect to the Bahamas Lithoherms. Site profiled via. coring and seismic 

reflections (Mullins and Neumann 1979). 

Santaren 

Mounds

Deep water mounds are described at 500-800 m along the western shelf edge of the Great 

Bahama Bank, running from the Santaren Channel to the Straits of Florida. Mounds >30 m 

were reported. Site profiled via. multi-beam side scan sonar and drop camera (Correa et 

al. 2006; Grasmueck et al. 2006).

Santa Marta 

Bank

This site is described as a deep water coral bank on the northwestern shelf of Colombia 

at a depth of 200 m. The bank is dominated by the potential structure-forming deep water 

coral M. myriaster. Anomocora fecunda, Coenosmilia arubuscula, and Polymyces fragilis

were reported abundant. Black corals and octocorals were reported numerous and include 

Antipathes columnaris, Elatopathes abetina, Sticopathes spp., Chrysogorigia desbonni, 

and Nicella guadalupensis. Site profiled via echosounding and dredges (Reyes et al. 

2005).

San Bernardo 

Bank

This site is described as a deep water coral bank on the northern shelf of Colombia at a 

depth of 155-160 m. The bank is dominated by the potential structure-forming deep water 

coral M. myriaster. Other species considered to contribute to habitat include A. fecunda, 

C. arubuscula, Eguchipsammia cornucopia, M. oculata, Javania cailleti, Caryophyllia 

berteriana, P. fragilis, Thalamophyllia riisei, Oxysmilia rotundifolia, and other Madracis

species. Flounder, basslets, and scorpionfish were the most abundant species of fish 

recovered. Madracis thickets and coralline mounds (up to 10 m in height) are reported. Site 

profiled via. echosounding, dredges, and grab sampling (Reyes et al. 2005; Santodomingo 

et al. 2006).

Table 8.2. Scleractinian-dominated coral habitats (“reefs”) of the wider Caribbean. Locations are identified in 

Figure 8.3.

deep-water coral banks off Colombia (Reyes et al. 

2005; Santodomingo et al. 2006). Investigations 

with sonar profiling and trawl and grab sampling 

suggest deep water banks from two locations 

off Columbia’s continental shelf at depths of 

155 - 200 m; Santa Marta and San Bernardo 

Banks. Table 8.2 details these locations. Relief, 

“suggestive of reef structures”, was reported. M. 

myriaster occurs in the U.S Caribbean (Figure 

8.5). Reyes et al. (2005) also report a shallower 

azooxanthellate coral community off Colombia at 

a depth of 70 m. Cladocora debilis was noted as 

the main matrix builder of this community. As this 

community occurs at less than 100 m in depth, C. 

debilis is not considered further.

Two other potential structure-forming species, D. 

alternata and D. dianthus are found throughout 

the wider Caribbean, but have not been reported 

from the U.S. Caribbean. D. alternata is found 

throughout the North Atlantic and forms bushy 

colonies to 1 m in height. It has been described as 

a reef builder (Le Goff-Vitry 2003) and has been 

recorded with M. oculata and M. myriaster in the 

wider Caribbean. D. dianthus is reported to form 

densely branched colonies and to “contribute to 
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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318

Figure 8.12. Large deep water gorgonian 

(Paracalyptrophora sp.) and deep water sponge 

(Pachastrellidae sp.). Photographed from 

submersible at 2,193 m in The Bahamas (Straits 

of Florida) (gorgonian is approximately 53.5 cm in 

height). Photo credit: Reed 2006.

Figure 8.13. 

Bubblegum coral 

(Paragorgia johnsoni) 

sampled by submersible 

from 620 m west of 

the Little Bahama 

Bank in the area of 

Bahamas Lithoherms 

(approximately 51 cm 

in height). Photo credit: 

Amy Wolfrum.

Figure 8.14. Large deep water gorgonian 

(Paramuricea sp.) providing habitat for an 

unidentified crustacean. Photographed from 

submersible at 2631 m in The Bahamas (Straits of 

Florida off Cat Cay) (gorgonian is approximately 33 

cm in width). Photo credit: Reed 2006.

Figure 8.15. 

Bamboo coral 

(Keratoisis flexibilis) 

on deck and utilizing 

a deep water 

structure-forming 

coral (E. profunda) 

as substrate 

(bamboo coral 

is approximately 

34 cm in height). 

Coral recovered 

from 806.5 m by 

submersible. Photo 

credit: Reed 2006.

Figure 8.11. Large 

deep-sea black 

coral (Leiopathes 

glaberrima) 

recovered from 

304 m in the 

North Atlantic 

(approximately 116 

cm in height).
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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319

reef formation” (Cairns and Stanley 1982). D. 

alternata and D. dianthus are detailed in Table 

8.2.

The distribution of major and potential structureforming species in the study area generally 

follows the Antillian arc and the margins of the 

continental shelves of South and Central America 

(Figure 8.3).

b. Black corals (Class Anthozoa, Order 

 Antipatharia, Families Cladopathidae and 

 Schizopathidae)

Approximately thirty-two species of antipatharia 

are reported from the wider Caribbean in depths 

below 100 m, with thirteen considered potentially 

important habitat-forming species. Eight species 

are reported from the U.S. Caribbean, with five 

considered potentially important habitat-forming 

species: Antipathes americana, A. caribbeana, 

Plumapathes pennacea, Tanacetipathes hirta, 

Parantipathes tetrasticha. Opresko and Sanchez 

(2005) list two regional black coral species of 

commercial importance: A. atlantica and A. 

gracilis. Both species are considered potentially 

habitat-forming. The maximum reported height 

of deep water antipatharians occurring in U.S. 

Caribbean waters is 61 cm (P. tetrasticha). Figure 

8.11 illustrates a large deep-sea black coral. 

c. Gold corals (Class Anthozoa, Order 

 Zoanthidea)

One species of gold coral is reported from the 

Caribbean region. Messing et al. (1990) describe 

a large zoanthidean tentatively identified as 

Gerardia sp. in Bahamian waters. This species 

is considered habitat-forming as colonies 1 m tall 

and 1.5 m across were reported. Gerardia sp. 

was found in dense stands on lithoherm crests, 

where current flow was the greatest.

d. Gorgonians (Class Anthozoa, Order 

 Gorgonacea) 

Approximately one hundred and forty-seven 

species of gorgonians are reported from the 

wider Caribbean at depths below 100 m. Thirteen 

species are reported from the U.S. Caribbean 

(more are expected). Forty-four species are 

considered potential habitat-forming species, 

with twelve occurring in U.S. Caribbean waters: 

Acanthogorgia goesi, Callogorgia americana 

Americana, Diodogorgia nodulifera, Ellisella 

barbadensis, E. elongata, Narella bellissima, 

N. pauciflora, N. deichmannae, N. obesa, N. 

guadelupensis, Riisea paniculata and Swiftia 

exserta. The maximum reported height of a 

wider Caribbean deep water gorgonian is 300 cm 

(Pseudoplexaura porosa), and 244 cm for U.S. 

Caribbean waters (E. barbadensis). Two species 

of red or pink corals (family Coralliidae); 13 species 

of bamboo coral (family Isididae); one species 

of bubblegum coral (family Paragorgiidae); and 

34 species of red tree coral (family Primnoidae) 

are known from the region. Detailed distribution 

data (beyond occurring in the western Atlantic) 

was available for only 54% (82) of known deep 

water gorgonians. Height and/or width data 

was available for 34% (50) of known deep water 

gorgonians. It is expected that many more 

gorgonians are potential habitat-forming species. 

Figures 8.12 - 8.16 and 8.17 illustrate deep-sea 

gorgonians.

Figure 8.16 Large deep water gorgonian Nicella

americana, collected from 165 m off the Dominican 

Republic (approximately 53 cm in width).

Figure 8.17. Deep water coral habitat at 829.4 m in 

The Bahamas (Straits of Florida). A variety of deepsea corals are illustrated, including the soft corals 

Anthomastus agassizi (red in color) and Pseudodrifa 

nigra (brown in color). Photo credit: Reed (2006).
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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320

Figure 8.18. Sea pens (Renilla reniformis), from 

Curacao, Netherlands Antilles (each approximately 2 

cm in width). This species occurs to 108 m or more.

e. True soft corals (Class Anthozoa, Order 

 Alcyonacea and Suborder Stolonifera)

Twenty species of true soft corals (Alcyonacea) 

are reported from the wider Caribbean in 

depths below 100 m. Only one is considered 

a potentially important habitat-forming species; 

Neospongodes portoricensis, with a maximum 

reported height of 30 cm. Only one is reported 

from the U.S. Caribbean: Stereotelesto corallina. 

Detailed distribution data was available for 35% 

(7) of known deep water alcyonaceans. Figure 

8.17 illustrates a deep-sea soft coral.

f. Pennatulaceans (Class Anthozoa, Order

 Pennatulacea)

Pennatulaceans, or sea pens, live in softsediment and are known to form extensive 

groves in some areas (Morgan et al. 2006). Eight 

deep-water sea pens are reported from the wider 

Caribbean. One species is reported from the U.S. 

Caribbean: Renilla reniformis, with a maximum 

reported height of 6.5 cm it is not considered an 

important habitat-forming species (illustrated in 

Figure 8.18). It is bioluminescent. The maximum 

reported height of a deep water pennatulacean 

occurring in the wider Caribbean is 210 cm, for 

Funiculina quadrangularis. According to Picton 

and Howsen (2002) F. quadrangularis is often 

found in very large colonies. This order contains 

the deepest deep-sea corals, many occur below 

1,000 m, with one species (Umbellula magniflora) 

reported from 6,300 m in the Caribbean Sea 

basin. Most deep water sea pens are found on 

basin floors, rises and trenches throughout the 

Caribbean region including the Caribbean Sea 

basin, Aves Ridge and Puerto Rico and Cayman 

Trenches. Height and width data were available 

for 25% (2) of known deep water sea pens.

g. Stylasterids (Class Hydrozoa, Order 

 Anthoathecatae, family Stylasteridae)

Forty-one species of stylasterids are reported 

from the wider Caribbean at depths below 100 

m. Only one is regarded as a potential habitatforming species; Distichopora sulcata (Figure 

8.19), with a maximum reported height of 25 

cm and width of 30 cm, endemic to the waters 

off northwest Cuba. However, additional larger 

stylasterids are likely - during a submersible dive 

James and Ginsburg (1979) recorded a larger 

unidentified stylasterid measuring “about 30 cm 

across” growing out from a rock wall at 290 m 

off Belize Glovers Reef). Fifteen smaller (non 

habitat-forming) species (<25 cm in height or 

Figure 8.19. Large deep water stylasterid 

(Distichopora sulcata) dredged off Havana, Cuba 

(approximately 33 cm in width).

Figure 8.20. Deep-sea stylasterid (Stylaster 

miniatus) providing substrate for a solitary deep 

water stony coral (Paracyathus pulchellus) and other 

unidentified invertebrates, the Straits of Florida, at 

191 m (stylasterid is approximately 10 cm in width).
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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321

width) occur in U.S. Caribbean waters. All wider 

Caribbean stylasterids are referenced by Cairns 

(1986). 

Puerto Rican waters contain two stylasterid 

species: S. atlanticus and S. spatula. Stylasterids 

have been recorded with many other species of 

deep water corals and can serve as substrate 

allowing invertebrates to gain a foothold (Figure 

8.20).

Deep Coral Habitat Types

Deep sea corals often occur as individual and/

or scattered colonies on the sea floor (Cairns 

1979, 2000). More numerous colonies can form 

clumps or thickets of varying density (Mullins et 

al. 1981; Messing et al. 1990) (Figures 8.7, 8.8 

and 8.10). These bush-like aggregations can 

form habitat on suitable substrate and contribute 

to the formation of deep sea bio-buildups 

through their skeletal debris and by trapping 

pelagic sediment. Two types of deep-sea biobuildups are reported from the wider Caribbean 

region (deep coral mounds and lithoherms); their 

morphologies and compositions, and deep-sea 

coral banks are discussed below. Seamounts 

are also discussed.

Deep Coral Mounds

A variety of terms have been used for the 

unconsolidated sea floor elevations with 

which deep-sea corals occur. In general these 

structures are described as “capping mounds of 

unconsolidated sediment and coral rubble, which 

are often built on an underlying lithified substrate” 

Figure 8.21. Idealized regional deep coral mound (not to scale). Associated benthos as follows: 1) Etmopterus 

sp. (dogfish sharks); 2) pteropods; 3) Brotulid sp. (fangtooths); 4) Diretmus sp. (dorys); 5) Myctophid sp. 

(lanternfish); 6) Macrouid sp. (grenadiers); 7) Stomias sp. (dragonfish); 8) Hexactinelida sp.; 9) Cirripathes sp.; 

10) S. varibilis; 11) M. oculata; 12) Galatheid sp. (deep-sea crabs); 13) crinoids; 14) gorgonians; 15) Iridogorgia

sp.; 16) Antipathes sp.; 17) Macruan sp.; 18) Keratoisis sp.; 19) Munida sp. (squat-lobsters); 20) Stylasteria

sp.; 21) Nematocarcinidae sp. (deep-sea shrimp); 22) Glyphocrangon sp.; 23) solitary corals; 24) Chrysogorgia

sp.; 25) Synaphobranchus sp.; 26) Lithodid sp.; 27) Nymphaster sp.; 28) porifera; 29) Alcyonaria sp.; 30) 

Nephropsis rosea; 31) Holothurian sp. 32) Ophiuroid sp.; 33) Turrida sp.; 34) Cruriraja sp. (skates). Sources 

listed in Appendix 8.2.
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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322

(Reed 2002). These structures have been called 

deep-water coral banks; deep-sea coral mounds; 

colony thickets; coral thickets; coral structures, 

deep-water coral reefs; and bioherms (Stetson 

et al. 1962; Squires 1964; Neumann et al. 1977; 

Mullins et al. 1981; Reed 2002). 

The accepted geological term for 

mounds rich in unbound skeletal 

debris is biostrome (Bates and 

Jackson 1987). In what follows 

we use “deep coral mounds” for 

this type of buildup. 

In the study region these deep 

coral mounds occur in areas 

with strong currents and/or 

upwelling where they accrete by 

a combination of coral skeletal 

debris and trapped suspended 

sediment swept into coral 

structures (Mullins et al. 1981). 

The resulting structures range in 

height over the surrounding sea 

floor from a few meters to 50 m 

(Reed 2002).

The most extensive known occurrence of deep 

coral mounds reported in the Caribbean is found 

on the northern slope of Little Bahama Bank at 

depths between 1,000 to 1,300 m (Mullins et 

al. 1981). Here, where bottom currents were 

reported to be as much as 50 cm/sec and 

temperatures of 4-6o C, numerous banks are 

inferred to occur in an area of 2,500 km2 with 

relief over the surrounding sea floor ranging 

from 5-40 m (ibid). The surface sediments of 

these mounds consist predominantly of pelagic 

organisms such as planktonic foraminifera and 

pteropods, with small amounts of coral skeletal 

debris and other invertebrates in the coarser 

grain sizes (ibid). These mounds appear to be 

developed by the baffling action of the thickets 

of deep corals that traps pelagic sediments 

and build on unconsolidated mounds. Other 

coelenterates found on these mounds include 

gorgonians, alcyonaceans and antipatharians 

(Figure 8.21). Mullins et al. (ibid) present a useful 

schematic model of the sequential development 

of these mounds (Figure 8.22) based on previous 

publications (Squires 1964; Neumann et al. 1977). 

Figure 8.3 illustrates the distribution of deep coral 

mounds. Table 8.2 details these occurrences.

Lithoherms

In contrast to the unconsolidated deep coral 

mounds, lithoherms (Neumann et al. 1977; 

Messing et al. 1990; Reed 2002) are composed 

of varied communities of benthic organisms 

growing on successive crust-like layers of friable 

limestone. Lithoherms have also been referred 

Figure 8.23. Idealized lithoherm of the Straits of Florida (not to scale). 

Figure adapted from Messing et al. 1990.

Figure 8.22. Hypothetical development sequence of 

deep coral mounds (not to scale). Figure adapted 

from Mullins et al. 1981, based in part on Squires 

1964 and Neumann et al. 1977.
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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323

to as bioherms, deep-water biohermal buildups 

and muddy carbonate buildups (Neumann et 

al. 1977; Reed 2002). Regionally, they are 

reported only from the eastern side of the Straits 

of Florida at depths between 500 and 700 m 

below sea level. According to Messing et al. 

(1990) the lithoherms are elongate mounds up to 

300 m long and 50 m high. Their surfaces show 

successive friable limestone layers composed 

of fine-grained carbonate skeletal sediment with 

variable amounts of coral debris. The cement of 

the crust-like layers of lithoherms is composed of 

magnesium calcite with 10-14 mole % magnesium 

(Wilbur and Neumann 1993). 

Messing et al. (1990) describe and illustrate (Figure 

8.23) a distinct, tripartite biozonation of one of 

these lithoherms, which they ascribe to variations 

in current velocity produced by the mounds, also 

described by Genin et al. (1986). On the steeper 

margin or side facing into the current (upcurrent 

margin) where velocities may reach 100 cm/

sec, corals (L. pertusa) predominate (“corals” in 

Figure 8.23). A zone of zoanthids forms along 

the upcurrent crest (gold coral = “zoanthids” in 

Figure 8.23) and crinoids and alcyonarians occur 

behind this zone (“crinoids and alcyonarians” in 

Figure 8.23) and on the flanks of the lithoherms 

(Genin et al. 1986). Figure 8.3 illustrates the 

distribution of known lithoherms. Table 8.2 details 

these occurrences.

Deep-sea Coral Banks

Reyes et al. (2005) describes two azooxanthellate 

deep water coral habitats (banks) at depths 

greater than 100 m off Colombia’s Caribbean 

shelf. Substrate included hard bottom limestone 

(fossilized reef) and metamorphic rocks. It is 

unclear to what extent these corals are building 

upon themselves (i.e. as with bio-buildups) or 

simply opportunistically taking advantage of 

suitable substrate (limestone platform). Reyes et 

al. (ibid) suggested similarities with shallow water 

reefs and stated a need for further investigations. 

Figure 8.3 illustrates the distribution of known 

deep water coral banks. Table 8.2 details these 

occurrences.

Seamounts

Deep sea corals are known as intimate associates 

of seamounts throughout the world (Freiwald et 

al. 2004; Clark et al. 2007). EarthRef.org (2007) 

lists four seamounts occurring within the U.S. 

Caribbean waters: Grappler Seamount; Whiting 

Seamount; Mona Seamount; and Dos Ni’os Knoll. 

Figure 8.5 illustrates their locations. The potential 

that seamounts throughout the region are home 

to deep water coral ecosystems deserves further 

attention.

Relevant Geological and Oceanographic 

Processes

Five processes play significant roles in the 

distribution and composition of bio-buildups: (1) 

foremost is amplification of current velocities 

by submarine topography; (2) the biological 

production of skeletal debris; (3) physical trapping 

and stabilization of sediment by thickets of corals, 

sponges and alcyonarians to build elevated 

accumulations; (3) availability of hard substrate 

for recruitment of corals; and (4) the submarine 

cementation that produces lithoherms. Process 3 

is the most significant for deep water banks.

Increased current velocities appear essential for 

flourishing deep corals both because they can 

locally sweep away sediment to expose the hard 

substrate that encourages settlement of coral 

larvae and they provide a supply of plankton 

on which the corals feed. In addition, increased 

currents bring a continued supply of pelagic 

sediment to help build local relief (mounds and 

lithoherms). 

Two kinds of submarine topography can 

accelerate current velocities. One, where water 

flow meets or is constricted between land masses, 

for example, the seaways or channels between 

islands of the Lesser Antilles or Windward 

Islands (Figure 8.2). The larger and deeper inter 

island spaces produce the major seaways; the 

more numerous, smaller and shallower ones 

produce similar but smaller increases in velocity. 

A second kind of topographic constriction occurs 

when water flow meets elevations of the sea 

floor, typically platforms or banks of calcareous 

deposits. The spaces between large and longlived platforms of the Bahamian Archipelago and 

the Florida Peninsula are sites of increased flow 

velocities between adjacent platforms. A smaller 

scale example of topographic constriction of 

currents is present in the series of shallow banks 

of calcareous deposits, which have developed 

on the Nicaragua rise and channel flow from the 

Caribbean Sea contributing to the initiation of 

the swift Yucatan Current. Localized sediment 

accumulation, either physical or as a result of 
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

324

the trapping by thickets of erect invertebrates, is 

needed to produce sea floor relief. 

The role of hard substrate in initiating coral 

settlement is ambiguous. On the one hand it is 

repeatedly cited as necessary for settlement of 

coral larvae (Freiwald et al. 2004, pp. 18, 21). 

However, in the same publication (ibid, pp 27-

28) occurrences are cited of deep-sea corals on 

mounds, which presumably are unconsolidated 

sediment. Mullins et al. (1981) described and 

illustrated a compelling example of corals growing 

on mounds of unconsolidated pelagic sediment 

from the margin of Little Bahama Banks. One 

explanation for this apparent contradiction may 

be the minimal size of hard substrate required to 

promote settlement. If for example, something 

as small as a bivalve shell is sufficient then 

deep-sea corals could establish on mounds of 

unconsolidated sediments that included bivalve 

skeletons. Once a few corals are established 

their continued production of skeletal debris 

provides continually expanding hard substrate 

for larval settlement (Figures 8.15 and 8.20 

illustrate the substrate use of deep-sea corals). 

Syndepositional, subsea cementation triggered 

perhaps by persistent strong currents (Straits of 

Florida) is required to produce current-resistant 

lithoherms and it may also be responsible for 

developing hard substrates on which deep coral 

mounds develop.

V. SPATIAL DISTRIBUTION OF CORAL 

 SPECIES AND HABITAT

The distribution of deep water corals generally 

follows the Antilles and continental shelves of 

Central, South and North America (Figures 8.3 - 

8.5). Dawson (2002) analyzed the distribution of 

129 species of azooxanthellate (deep-sea) stony 

corals in the Caribbean and surrounding areas. 

His findings include the following: the greatest 

diversity of species occurs at around 200-350 

m in depth; and the highest diversity of species 

is located at the northern islands of the Lesser 

Antilles.

To date, no significant deep scleractinian coral 

habitats have been discovered in the U.S. 

Caribbean. Deep-sea corals have been reported 

from submersible and ROV research, but only 

sporadically. However, deep-sea research for 

the region is still in its infancy. Previous research 

includes limited ROV and submersible work off 

Navassa Island (Littler and Littler 1999, Reed 

pers. comm.) and limited submersible work off 

Puerto Rico and the U.S. Virgin Islands (Nelson 

and Appeldoorn 1985). Figure 8.24 illustrates 

a cross section of a “typical” continental slope 

for Puerto Rico and the U.S. Virgin Islands, 

adapted from Nelson and Appeldoorn (1985). 

Intermittent zones of rubble, rocks and barren 

sand with occasional sponges, hydroids, soft 

corals, hard corals, crinoids and gorgonians 

attached to rocky substrate are described. 

“Sea whip” type gorgonians (Ellisella sp.), often 

wrapped with basket stars, were noted common 

at one location off the south coast of Puerto Rico. 

Rugged rock habitat, described as large, closelypacked boulders with little or no sand, was noted 

to primarily occur in the Mona Passage. Most 

notably, an abundance of the potential structureforming coral M. myriaster was reported at a 

depth of 197-212 m on the shelf margin south of 

the southwest corner of Puerto Rico.

Ongoing deep water research in the U.S. 

Caribbean includes the exploration and 

characterization of seafloor habitats, with 

ROVs, down to 1,000 meters within the U.S. 

Virgin Islands and Puerto Rico by NOAA (NOAA 

2007). This effort aims to characterize and map 

moderate to deep water coral reef ecosystem 

habitats within targeted study areas. Priority 

areas include the deep waters of the Buck Island 

Reef National Monument, the Escollo Grappler 

seamount to the southeast of Puerto Rico and 

Figure 8.24. Cross-section of a “typical” continental 

slope for Puerto Rico and the U.S. Virgin Islands to 

approximately 400 m. Deep-sea research has been 

limited and, to date, deep corals have been only 

sporadically reported from the region. Figure adapted 

from Nelson and Appeldoorn 1985.
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

325

the La Parguera region along the southwestern 

coast of Puerto Rico.

Inspection of regional unpublished research 

reports and dredge station sorting sheets 

illuminate interesting possibilities for deep coral 

habitat. Selected citations from dredge reports 

(individual station records) and notes include the 

following:

For the U.S. Caribbean: in Navassa Island waters 

two major structure forming corals (M. oculata and 

E. rostrata) were recovered, at 1,200 m, along with 

solitary corals (Stephanocyathus laevifundus and 

of the families Deltocyathus and Desmophyllum), 

gorgonians (of the genus Acanella, Iridogorgia

and Chrysogorigia) and deep water fish (Staiger 

and Voss 1970).

For waters adjacent to the U.S. Caribbean: in 

British Virgin Islands waters, at 205 - 380 m, 

one potential structure-forming coral species 

(M. myriaster) was recovered from the same 

location in a trawl net “festooned with stalked 

crinoids”. Deep water fish and crustaceans were 

also recovered (Staiger 1969); in Saba waters, in 

512 m off the northern edge of the Saba Bank, 

a large deep water gorgonian (C. multiflora) was 

recovered along with deep water fishes (Anon. 

1956); south of Hispaniola in 135-169 m, the 

potential structure forming coral species (M. 

myriaster) was recovered along with numerous 

sponges, gorgonians and other invertebrates 

in a net noted as “We had to use the crane to 

lift it aboard.... The catch was very good”. Deep 

water fish and crustaceans were also recovered. 

Similar “excellent” and “magnificent” catches were 

also noted from two nearby trawl sets that also 

recovered M. myriaster (Staiger and Voss 1970; 

Voss 1971).

For waters throughout the region: in Venezuelan 

waters with L. pertusa at 406 m - “it was the best 

10 ft. otter trawl tow I have seen in six years. 

There was a greater variety of organisms and a 

greater number than we usually get in a 41 ft. otter 

trawl tow” (Anon 1968); in the Straits of Florida, 

in waters just off just off Havana, Cuba, where M. 

carolina, M. oculata, and M. myriaster have been 

recovered - “the “pentacrinus grounds” [crinoid 

grounds] discovered by the Blake off Havana are 

world famous” (Voss 1966b); and in the Straits of 

Florida just southeast of the area of lithoherms on 

the Little Bahama Bank) with L. pertusa, S. varibilis

and E. profunda at 738 m - “Tremendous amount 

of coral in net. Needed crane to lift it aboard. Good 

catch of fish and invertebrates” (Anon. 1973).

Though no direct information was found 

regarding the regional transport of deep coral 

larvae, the cosmopolitan distribution of shallowwater benthos of coral reefs as well as many 

deep water coral species (including the major 

structure-forming corals) indicates long term 

connectivity throughout the basin. 

VI. SPECIES ASSOCIATIONS WITH DEEP

 CORAL COMMUNITIES 

Fauna of deep-sea bio-buildups are among 

the least studied of marine assemblages. 

For example, the identification of deep-sea 

gorgonians, collected in the Caribbean and the 

Bahamas in the early 1960’s, are, to date, being 

revised (e.g., Cairns 2007); deep-sea Gold corals 

collected on lithoherms in the Bahamas in the 

1970’s still await full identification (Messing pers. 

comm.), and crustaceans collected in the 1880’s 

have long waited proper scientific descriptions 

(Larsen 1999).

Inspection of archival data and grey literature, 

including original station data sheets, cruise 

narratives and biological survey reports from 

28 locations within the region identify a wide 

assortment of associated species occurring with 

the major structure-building deep-water corals; E. 

profunda, L. pertusa, M. carolina, M. oculata and 

S. varibilis (Appendix 8.2). Figure 8.21 illustrates 

several of these species in an idealized deep 

coral mound. Many of the reported associated 

species may grow to almost one meter in height 

(larger cnidarians and porifera).

Large deep sea sponges (phylum Porifera) are 

a potentially important megafauna component of 

deep sea coral habitats. Deep sea sponges have 

been recorded with structure-forming and potential 

habitat-forming corals (Appendix 8.2) and on 

deep water mounds, lithoherms and banks. It is 

unknown what habitat role they may play on deep 

water coral habitats. Colin (1978) reports that in 

some areas of shallow water reefs the biomass 

of sponges “can exceed that of any other group, 

including reef-building corals.” Lang et al. (1975) 

reported that sponges (sclerosponges) can act 

as primary framework builders of reefs at depths 

of 70-105 m off Jamaica. Of primary interest are 

the larger calcareous sponges (class Calcarea), 

deep water glass sponges (class Hexatinellida), 

demosponges (class Demospongiae) and 

sclerosponges (class Sclerospongiae). Deep-sea 

sponges have been recorded up to 122 cm wide 
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

326

Measure of Impact

Gear Type Current Fishery 

Use in Region

Potential 

Severity of 

Impact

Potential Extent 

of Impact from 

Fishing Gear

Current 

Geographic 

Extent of Use 

in Region

Overlap 

of use 

with coral 

habitat

Overall 

Rating 

of Gear 

Impact

Bottom Trawl None reported High High NA Uncertain NA

Mid-water Trawl None reported Low Low NA Uncertain NA

Dredge None reported High Low NA Uncertain NA

Bottom-set Longline to 109 m Medium Low Unknown Uncertain Low

Bottom-set Gillnet None reported Medium Medium Unknown Uncertain Low

Longline traps or pots to 183 m High Medium To shelf edge Uncertain Low

Single pots to 183 m Low Medium To shelf edge Uncertain Low

Hook and line to 600 m Low Low Unknown Uncertain Low

Table 8.3. Potential fishing gear impacts on deep sea corals for the U.S. Caribbean.

at deep-sea coral depths in the Straits of Florida. 

Figure 8.12 illustrates a deep-sea sponge. Deepsea sponges are subject of pharmacological 

investigations.

A number of fishes have been observed or 

collected in association with deep coral habitats. 

While most of these are demersal species 

(Appendix 8.2), certain commercially important 

pelagic species may also associate with deepsea coral habitat. Swordfish (Xiphias gladius) 

have been encountered by submersibles 

visiting deep sea corals (Lophelia) in the Straits 

of Florida (Nizinski and Ross 2002) and in 

Bahamian deep waters at 600 m (Harbison and 

Janssen 1987). Larger deep coral mounds and 

lithoherms may play similar roles to seamounts, 

which are thought to attract pelagic fish because 

of an “enhanced primary productivity due to 

particular oceanographic conditions supporting 

a rich ecosystem” (Allain et al. 2006). In effect, 

seamounts influence oceanic currents in a way 

that promotes plankton to develop, acting as a 

feeding source for resident fish, attracting larger 

pelagic predators. These pelagic species are 

thought to use the seamounts unique underwater 

features primarily for food but also potentially as 

spawning grounds and/or nursery areas. 

VII. STRESSORS ON DEEP CORAL 

COMMUNITIES

Serious potential threats to deep water coral 

communities found in other U.S. waters are 

not currently found in the U.S. Caribbean. No 

current examples were found of significant 

threats to deep water corals from the following: 

harvesting of precious coral, cable laying, oil and 

gas exploration, sedimentation, invasive species 

and destructive fishing practices (such as bottom 

trawling). 

Fishing occurs to deep coral depths (>100 m) in 

the U.S. Caribbean and throughout the region. 

This activity is mainly focused on deep water 

snapper and grouper and includes commercial 

and recreational fishing effort. An extensive 

deep water fishery, primarily for snappers, occurs 

around Puerto Rico (Appeldoorn pers. comm.). 

Target species include black snapper (Lutjanus 

griseus), blackfin snapper (L. buccanella), 

vermilion snapper (Rhomboplites aurorubens) 

and silk snapper (L. vivanus) (FAO 1993; Kojis 

2004; Cummings and Matos-Caraballo 2003). 

Gear utilized in commercial deep water snapper 

and grouper fishing in the U.S. Caribbean includes 

vertical set line, bottom longline, handlines, 

electric or hydraulic reels and traps (Kojis 2004; 

Tobias 2004). Similar effort is reported throughout 

the region (FAO 1993). Figure 8.25 illustrates 

a typical commercial fishing vessel capable 

of fishing offshore for deepwater snapper and 

grouper in the U.S. Virgin Islands. The maximum 

reported depth for line fishing is 366 m (Swingle 

et al. 1970). The maximum reported depth for 

trap fishing is 183 m (Sheridan et al. 2006). 

During submersible observations, Nelson and 

Appeldoorn (1985) encountered ghost fishing 
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

327

traps at a depth of 121 m off the west end of St. 

Croix and a discarded longline at 236 m off the 

harbor entrance to San Juan, Puerto Rico. 

Recreational fishing for deep water species in 

the U.S. Caribbean and throughout the region is 

termed “deep drop” fishing (also referred to as 

cannonball fishing). Gear includes expensive 

electric reels and heavy sash weights (commonly 

weighing 5 to 10 lbs) which allow fishing to 

potentially >600 m (Sword Fishing Central 2006). 

The Melton International Tackle Catalog (2007) 

lists electric deep drop reels up to $4,500 in 

price. Reels “capable of landing fish as deep 

as 500 fathoms [914 m]” are advertised (Melton 

2007, pg 95). Heavy weights are needed for 

optimal deep drop fishing performance; to take 

the line straight down to the sea floor and keep 

the tackle as close to the bottom as possible, in 

a process called “bouncing,” where the weight 

bounces up off the sea floor and the whole rig 

remains under the boat (Brooks 2007). Nonconventional weights, fashioned from pipes or 

other containers filled with concrete are also 

used. Deep drop fishing has recently gained 

in popularity with sport fishers, the 3nd annual 

‘Lords of the Deep’ Fishing Tournament was held 

in September 2007 in Nassau, Bahamas. Figure 

26 illustrates electric fishing reels utilized in deepdrop fishing. Such gear is available in the U.S. 

Caribbean. A small level of effort of deep-drop 

fishing is reported from the U.S. Caribbean and 

throughout the Caribbean region.

The association between deep-sea fishing and 

deep-sea coral habitat is unknown. However, a 

review of web-based fishing forums reveals that 

recreational sport fishers may be aware of and 

fishing on such: 

“You want to look for mounds, ridges, humps, 

etc. The key is learning the currents and exact 

location of your drop especially in very deep 

water (1000-2000 ft) [305-610 m]. You will find 

that on one particular mound, the peaks will 

hold one type of fish and the base of that same 

mound will hold another. Some deep water 

mounds in the Bahamas go up to a couple of 

hundred ft. [61 m] from the base. The Pomfret 

is one of my deep water favorites. We catch 

this species on the peaks of mounds. If your 

drop is off the mounds peak, you will have a 

tough time catching them.” (Offshore Fishing 

Forum 2006). 

Table 8.3 details potential fishing gear impacts on 

deep-sea corals for the U.S. Caribbean with deep 

drop fishing included under “hook and line”.

When compared to potential threats deep coral 

habitats experience in other U.S. waters (chiefly 

destructive bottom trawling) this effort does not 

currently pose a significant threat to deep corals. 

However, the potential for damage to deep corals 

from such fishing gear is well documented, and 

may increase as the amount of fishing effort 

increases (Chuenpagdee et al. 2003). Traps 

and weighted lines can crush structure-forming 

corals and entangle or snare softer octocorals. 

The potential impacts of regional commercial 

and recreational deep-sea fishing warrants 

further investigation. Additionally, the impact 

that the removal of large deep water 

predators may have on deep coral 

ecosystems is unknown. However, it 

is well documented that the overfishing 

of these species can have negative 

effects on shallow reef health (Dulvy 

et al. 2004; Steneck and Sala 2005; 

Mumby et al. 2006).

Figure 8.25. Bandit rigged boat 

- a type of commercial fishing 

vessel used to fish offshore for 

deepwater snapper and grouper 

in the U.S. Virgin Islands and 

capable of fishing to depths 

where deep water coral reside. 

Photo credit: Kojis 2005.

Figure 8.26. Typical electric reels used in deep drop 

fishing. This gear is marketed in the U.S. Caribbean. 
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

328

Current efforts to manage fisheries the Caribbean 

waters are exacerbated by a regional ‘lack of 

capacity’ in fisheries management (Haughton et al. 

2004). Traditional management strategies (such 

as catch and release) may not work for deep-water 

species, as fish brought up from deep-sea coral 

depths expire en-route. Additionally, remoteness 

of location may play an important role with regard 

the management of deep-sea coral habitats. 

For example, the major structure forming corals 

M. oculata and E. rostrata have been found in 

the waters surrounding Navassa Island (Figure 

8.5), an extremely remote uninhabited U.S. 

territory and illegal fishing activities are reported 

from Navassa’s waters (Miller 2003). However, 

the closest inhabited United States territory, 

Guantanamo Bay Naval Base (Cuba), is located 

approximately 170 km distant.

Throughout the region, future fishing activities 

pose a significant potential threat, as shallow 

water fisheries resources are depleted and 

commercial and recreational effort moves 

into deeper water (Prado and Drew 1999; 

FAO 2000; Koslow et al. 2000; Roberts 2002). 

Inspection of research station sorting sheets and 

current literature reveal numerous fish species 

associated with deep corals that belong to taxa 

currently harvested in other regions and may be 

harvested in the Caribbean in the future. Fishes 

include targeted and by-catch deep-sea species 

such as deep-sea codfish, rattails or grenadiers, 

hake, dory, sea robins, wreakfish (Polyprion 

americanus) and sharks (Merrett and Haedrich 

1997; Carpenter 2002; Freiwald 2004; Gordon 

2004) (locations of these sites are illustrated in 

Figure 8.3, species are listed in Appendix 8.2). 

Wreakfish (Sedberry et al. 1994) and dogfish 

sharks are currently harvested in U.S. continental 

waters. Wreckfish and have been observed on 

deep coral habitat (mounds and lithoherms) in the 

Straits of Florida (Messing et al. 1990). Dogfish 

sharks are reported on deep coral mounds 

and lithoherms (Nizinski and Ross 2002) and 

associated with occurrences of structure-forming 

corals throughout the region (Appendix 8.2). 

No significant threat of deep water trawling was 

found for the wider Caribbean. However, deep 

water shrimp trawlers are currently exploring 

fishing beyond 70 m in depth off Colombia (Gracia 

pers. comm.) and Reyes et al. (2005) note the 

need to asses the impacts of trawl fishing on the 

deep water coral banks off Colombia. 

Oil and gas exploration does currently occur in the 

to U.S. Caribbean. However, this activity occurs 

in the southeast Caribbean, notably off Trinidad 

and Tobago, and in Straits of Florida waters off 

northwest Cuba. It is unknown what effect these 

activities have on deep coral communities. It is 

also unknown to what extent regional underwater 

cables and lines affect deep coral communities. 

The harvest of precious corals, red and black 

corals, is prohibited in the U.S. Caribbean and 

there are no reports of current harvest activities. 

However, black coral jewelry, reportedly 

manufactured from corals harvested in the 

Cayman Islands, is available for purchase in the 

U.S. Virgin islands (Greenberg and Greenberg 

2006; Bernard K. Passman Gallery, St. Thomas, 

pers. comm.). Black coral sold in the U.S. 

Caribbean is reportedly harvested in the western 

Caribbean from coral beds located at depths 

of 61-122 m (200-400 ft) (Bernard K. Passman 

Gallery, St. Thomas, pers. comm.). Regional 

countries listed by the Convention on International 

Trade in Endangered Species of Wild Fauna and 

Flora (CITES) trade database as exporting black 

coral (Antipatharia spp.) since 2001 include the 

Bahamas, British Virgin Islands, Cayman Islands, 

Cuba, Colombia, Dominican Republic, Mexico, 

Panama, and Trinidad and Tobago (Figure 8.4, 

some locations may be transshipment points) 

(CITES 2007). The basil axis of black coral is 

used in the manufacture of jewelry (Opresko and 

Sanchez 2005).

VIII. CURRENT CONSERVATION AND 

MANAGEMENT ACTIVITIES 

In the U.S. Caribbean deep-water corals 

and other marine resources are under the 

jurisdiction of territory (USVI: 0-3 nmi from 

shore), commonwealth (Puerto Rico: 0-12 nmi 

from shore) and federal management authorities 

(U.S. EEZ and the waters around Navassa 

Island). Fisheries in the U.S. EEZ around USVI 

and Puerto Rico are managed under fishery 

management plans developed by the Caribbean 

Fishery Management Council (CFMC), based 

in Puerto Rico. Deep sea coral language in the 

recently reauthorized Magnuson-Stevens Act 

includes an affirmation of the Council’s authority 

designate zones to protect deep sea corals from 

physical damage from fishing gear as part of their 

fishery management plans, without having to 
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

329

prove that corals constitute essential fish habitat 

[as amended by the Magnuson-Stevens Fishery 

Conservation and Management Reauthorization 

Act (Public Law 109-479), see Chapter 1]. 

Navassa Island is managed by the U.S. Fish and 

Wildlife Service as a National Wildlife Refuge 

that includes a 12 mile radius of marine habitats 

around the island. Commonwealth and territorial 

management authorities include the Puerto 

Rico Department of Natural and Environmental 

Resources and the United States Virgin Islands 

Department of Planning and Natural Resources.

Current efforts to manage fisheries in Caribbean 

waters are focused on shallow-water coral reef 

areas and associated fisheries (conch, grouper, 

Caribbean spiny lobster, etc.) (Sala et al. 2001; 

Theile 2001; FAO 2003). However, the Territory 

of the U.S. Virgin Islands includes antipatharians 

(black corals) in its local Endangered Species 

Act (ESA), thus granting the authority to protect 

black corals (no matter the depth) within USVI 

territorial waters. Puerto Rico implements the 

Federal Endangered Species Act (Federal ESA) 

in local waters. However, currently, no deep 

coral species are listed under the Federal ESA. 

The Caribbean Council manages corals under 

the Corals and Reef Associated Invertebrates of 

Puerto Rico and the U.S. Virgin Islands Fishery 

Management Plan (Coral FMP) (Caribbean 

Fishery Management Council 1994). It lists many 

coral genera and species that are found in both 

shallow and deep waters. Those species found 

in deep water (including Iciligorgia schrammi

(deep sea fan), Telesto spp., Ellisella spp. (sea 

whips) and antipatharians (black corals) would 

be protected under the Coral FMP. There are no 

conservation or management efforts specifically 

targeting deep-water corals for the U.S. Caribbean 

or the wider Caribbean region.

An assessment of regional deep-sea recreational 

and commercial fishing effort, especially deep 

drop fishing, would provide managers with 

information concerning the potential impact 

of this activity has on deep-sea corals. The 

collection of socioeconomic information and 

perceptions and attitudes to policy options, along 

with standard fisheries information (such as 

fishing effort, frequency of activity, landing and 

bycatch information, etc.), could help minimize 

any negative impacts of policies directed at the 

conservation of deep sea coral ecosystems

IX. REGIONAL PRIORITIES TO 

 UNDERSTAND AND CONSERVE DEEP 

 CORAL COMMUNITIES

Morgan et al. (2006) advocate that ecosystembased management, research and mapping and 

a ban on destructive fishing practices in deep sea 

coral areas are necessary to provide sufficient 

protection for deep-sea coral ecosystems.

Ongoing projects that may generate information 

regarding the distribution deep coral communities 

include the previously mentioned benthic habitat 

characterization effort by NOAA (NOAA 2007) 

and a coral reef assessment and habitat mapping 

effort of Navassa Island National Wildlife Refuge 

by NOAA and the National Park Service (Miller 

pers. comm.). The Navassa Island effort, 

although currently focused on shallow water 

reefs, could include deep-water investigations. 

Currently, no activities in the U.S. Caribbean are 

focused on the biology and ecology of deep coral 

communities.

Mapping the locations of deep coral habitat 

would be a valuable component of any true 

ecosystem based management program for 

the U.S. Caribbean. Mapping the locations of 

coral habitats gives a crucial spatial framework 

for managers to better understand the effects 

of ocean uses on the environment. Figure 8.5 

illustrates existing managed areas in the U.S. 

Caribbean and surrounding waters that have 

potential deep water coral habitat (waters >100 

m). Navassa Island Wildlife Refuge is the only 

managed area that includes documented deep 

water structure-forming and potential habitatforming scleractinian corals (E. rostrata, M. 

oculata and M. myriaster).

Examples of archival data awaiting further 

inspection include the following: video footage 

and photographs recorded during the submersible 

observations around Puerto Rico and the U.S. 

Virgin Islands reported by Nelson and Appeldoorn 

(1985), stored at the National Marine Fisheries 

Service (NMFS) Southeast Fisheries Science 

Center in Pascagoula, Florida (Appeldoorn pers. 

comm.); video footage and dive records recorded 

during submersible observations at Navassa 

Island by the Center for Marine Conservation (now 

The Ocean Conservancy); station records from 

numerous research cruises kept on microfilm at 

the NMFS South East Fisheries Science Center 
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

330

in Miami, Florida; and cruise reports and other 

information kept at the academic institutions that 

carried out much original deep sea research 

- the University of Miami’s Marine Invertebrate 

Museum in Miami, Florida, is one such location. 

An assessment of regional deep-sea recreational 

and commercial fishing effort, especially deep drop 

fishing, would provide managers with information 

concerning the potential impact of this activity. 

The collection of socioeconomic information and 

perceptions and attitudes to policy options, along 

with standard fisheries information (such as 

fishing effort, frequency of activity, landing and 

bycatch information, etc.), could help minimize 

any negative impacts of policies directed at the 

conservation of deep sea coral ecosystems.

X. CONCLUSIONS

It is clear from this review that the U.S. Caribbean 

and the wider Caribbean region contain a 

huge diversity of deep water corals. However, 

information on regional distribution of deep corals 

is limited and needs great expansion if deep 

corals can be part of any meaningful ecosystem 

based management strategy. Nevertheless, the 

thorough description of deep coral mounds and 

lithoherms in the Bahamas and in the Straits 

of Florida and deep coral habitat off Colombia 

provides valuable information on the bathymetric 

and hydrographic conditions favoring the 

development of deep coral communities. From 

this information inferences may be made regarding 

the morphology, composition and organism 

zonation of regional deep coral communities and 

the potential for deep coral habitat in the U.S. 

Caribbean. 

The lithoherms of the Straits of Florida are 

remarkable examples of the interaction of 

strong currents with a varied suite of benthos 

(corals, zoanthids, crinoids, alcyonarians, etc.) 

a rain of pelagic sediments and syndepositional 

cementation of the sediments and skeletal 

debris. This interaction produces currentresistant structures with a continual supply of 

new hard substrate for an attached and boring 

benthic community. It is these same interrelated 

processes that allow the development of ocean 

facing, wave-resistant shallow reefs. There 

are strong indications that the Straits of Florida 

contain undiscovered large areas of deep coral 

mounds and/or lithoherms and banks. Similarly, 

it seems likely that other areas throughout the 

Caribbean, including United States territories, 

with stronger currents may also have deep coral 

ecosystems. The possibility that deep-sea coral 

habitat in these areas harbor fish, which are 

exploitable (Appendix 8.2 and 8.3), deserves 

further attention.

Technical note

All images by Steven J. Lutz unless otherwise 

noted. Images may be modified for clarification.
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

331

XI. REFERENCES

Agassiz A (1888) Three cruises of the Blake. 

Houghton, Mifflin, N.Y., 314 pp

Allain V (2006) Seamounts and Pelagic 

Fisheries Interactions Under Study. 

South Pacific Commission Fisheries 

Newsletter 116: 3 pp

Anon. (1956) Sorting sheets and dredge 

station notes from R/V Oregon, 

Station O 2636. NOAA, Southeast 

Fishery Science Center (library). 

Uncatalogued microfiche

Anon. (1963) Sorting sheets and dredge 

station notes from R/V Gerda, Station 

G 118. Marine Invertebrate Museum 

Rosenstiel School of Marine and 

Atmospheric Science (RSMAS) 

University of Miami

Anon. (1964) Sorting sheets and dredge 

station notes from R/V Gerda, Station 

G 251. Marine Invertebrate Museum 

RSMAS University of Miami

Anon. (1965) Sorting sheets and dredge 

station notes from R/V Gerda, 

Stations G 503, G 636, G 691, G 692. 

Marine Invertebrate Museum RSMAS 

University of Miami

Anon. (1966) Sorting sheets and dredge 

station notes from R/V Pillsbury, 

Stations P 364, P 388. Marine 

Invertebrate Museum RSMAS 

University of Miami

Anon. (1968) Sorting sheets and dredge 

notes from R/V Pillsbury, Sta P 607, 

P 636, P 741, P 747, P 755, P 776. 

Marine Invertebrate Museum RSMAS 

University of Miami

Anon. (1969) Sorting sheets and dredge notes 

from R/V Pillsbury, Stations P 881, P 

891, P 892, P 954. Marine Invertebrate 

Museum RSMAS University of Miami

Anon. (1970) Sorting sheets and dredge 

notes from R/V Pillsbury, Stations P 

1187 and P 1262. Marine Invertebrate 

Museum RSMAS University of Miami

Anon. (1972a) Sorting sheets and dredge 

station notes from R/V Gilliss, Station 

GS 31. Marine Invertebrate Museum 

RSMAS University of Miami

Anon. (1972b) Sorting sheets and dredge 

notes from R/V Columbus Iselin, 

Station CI 6. Marine Invertebrate 

Museum RSMAS University of Miami

Anon. (1973) Sorting sheets and dredge 

notes from R/V Columbus Iselin, 

Station CI 140. Marine Invertebrate 

Museum RSMAS University of Miami

Anon. (1974) Sorting sheets and dredge notes 

from R/V Columbus Iselin, Stations 

CI 148, CI 246. Marine Invertebrate 

Museum RSMAS University of Miami

Armstrong RA, Singh H; Torres J; Nemeth 

RS; Can A; Roman C; Eustice R; 

Riggs L; Garcia-Moliner G (2006) 

Characterizing the deep insular shelf 

coral reef habitat of the Hind Bank 

marine conservation district (US Virgin 

Islands) using the Seabed autonomous 

underwater vehicle. Continental Shelf 

Research 26: 194–205

Bates R, Jackson J (eds) (1987) Glossary 

of Geology. Third Edition, American 

Geologic Institute, Alexandria, Virginia: 

788 pp

Bayer FM (1952) New western Atlantic 

records of octocorals (Coelenterata: 

Anthozoa), with descriptions of 

three new species. Journal of the 

Washington Academy of Sciences 42: 

183-189

Bayer FM (1961) The Shallow Water 

Octocorallia of the West Indian Region. 

Studies on The Fauna of Curacao and 

other Caribbean Islands 12: 377 pp
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

332

Bayer FM (1966) Dredging and trawling 

records of R/V John Elliot Pillsbury 

for 1964 and 1965. Studies in Tropical 

Oceanography 4 (1): 82-105

Bayer FM (2001) New species of 

Calyptrophora (Coelenterata: 

Octocorallia: Primnoidae) from the 

western part of the Atlantic Ocean. 

Proceedings of the Biological Society 

of Washington 114 (2): 365-378

Bayer FM, Muzik KM (1977) An Atlantic 

Helioporacean coral (Coelenterata: 

Octocorallia). Proceedings of the 

Biological Society of Washington 90 

(4): 975-984.

Bayer FM, Muzik KM (1979) The correct 

name of the helioporan coral 

Lithotelesto micropora Bayer & Muzik. 

Proceedings of the Biological Society 

of Washington 92(4): 873-875

Bedford Institute of Oceanography (2003) 

Identification Sheets for Corals in 

Atlantic Canada. Marine Environmental 

Sciences Division, Bedford Institute 

of Oceanography, Department of 

Fisheries and Oceans, 8 pp

Brook G (1889) Zoology 80 LXXX: Report on 

the ANTIPATHARIA. By George Brook, 

F.L.S., F.R.S.E. Bound in Volume 32: 

222 pp

Brooks R (2007) Deep drop fishing. Bass Pro 

Shops, OutdoorSite Library. Undated 

world wide web article, accessed 

August 2007. Available on-line at: 

<http://www.basspro.com/webapp/

wcs/stores/servlet/CFPage?mode=ar

ticle&objectID=31353&storeId=10151

&catalogId=10001&langId=-1>

Cairns SD (1977) Guide to the commoner 

shallow-water gorgonians of South 

Florida, the Gulf of Mexico, and the 

Caribbean region. University of Miami 

Sea Grant Field Guide Series 6: 74 

pp

Cairns SD (1979) The deep-water scleractinian 

of the Caribbean Sea and adjacent 

waters. Studies on the Fauna of 

Curacao and other Caribbean Islands 

57: 1-341

Cairns SD (l986) A revision of the Northwest 

Atlantic Stylasteridae (Coelenterata: 

Hydrozoa). Smithsonian Contributions 

to Zoology 4l8: l3l pp

Cairns SD (1999) Species Richness of 

Recent Scleractinia. Atoll Research 

Bulletin 459: 13-46

Cairns SD (2000) A revision of the shallowwater azooxanthellate Scleractinian 

of the western Atlantic. Studies on the 

Natural History Caribbean Region 75: 

321 pp

Cairns SD (2001) Studies on western Atlantic 

Octocorallia (Coelenterata: Anthozoa). 

Part 1. The genus Chrysogorgia. 

Proceedings of the Biological Society 

of Washington 114(3): 746-787

Cairns SD (2005) Western Atlantic DeepWater (>200 m) Octocorallia. 

Unpublished list, dated 11/2005

Cairns SD (2007) Studies on western Atlantic 

Octocorallia (Gorgonacea: Elliseidae). 

Part 7: The genera Riisea Duchassaing 

& Michelotti, 1860 and Nicella Gray, 

1870. Proceedings of the Biological 

Society of Washington 120 (1): 1-38

Cairns SD, Bayer FM (2002) Studies 

on western Atlantic Octocorallia 

(Coelenterata: Anthozoa). Part 2: 

The genus Callogorgia Gray, 1858. 

Proceedings of the Biological Society 

of Washington 115(4): 840-867

Cairns SD, Bayer FM (2003) Studies 

on western Atlantic Octocorallia 

(Coelenterata: Anthozoa). Part 3: 

The genus Narella Gray, 1870. 

Proceedings of the Biological Society 

of Washington 116(2): 617-648
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

333

Cairns SD, Bayer FM (2004a) Studies 

on western Atlantic Octocorallia 

(Coelenterata: Anthozoa). Part 4: The 

genus Paracalyptrophora Kinoshita, 

1908. Proceedings of the Biological 

Society of Washington 117 (1): 114–

139

Cairns SD, Bayer FM (2004b) Studies 

on western Atlantic Octocorallia 

(Coelenterata: Anthozoa). Part 5: 

The genera Plumarella Gray, 1870; 

Acanthoprimnoa, n. gen.; and 

Candidella Bayer, 1954. Proceedings 

of the Biological Society of Washington 

117 (4): 447–487

Cairns SD, Bayer FM (2002) Studies 

on western Atlantic Octocorallia 

(Coelenterata: Anthozoa). Part 2: 

The genus Callogoria Grey, 1858. 

Proceedings of the Biological Society 

of Washington 115 (4): 840-867

Cairns SD, Bayer FM (2006) Studies 

on western Atlantic Octocorallia 

(Coelenterata: Anthozoa). Part 6. 

The genera Primnoella Gray, 1858; 

Thouarella Gray, 1870; Dasystenella

Versluys, 1906. Proceedings of the 

Biological Society of Washington 

119(2): 161-194

Cairns SD, Stanley G (1982) Ahermatypic coral 

banks: Living and fossil counterparts. 

Proceedings of the Fourth International 

Coral Reef Symposium, Manila (1981) 

1: 611-618

Cairns SD, Zibrowius H (1997) Cnidaria 

Anthozoa: Azooxanthellate 

Scleractinia from the Philippine and 

Indonesian regions. In: Crosnier In: Crosnier 

A, Bouchet P (eds) Resultats des 

campagnes MUSORSTOM, volume 

16. Memoires du Museum national 

d’histoire naturelle 172: 27-243

Caribbean Fishery Management Council 

(1994) Fishery Management Plan, 

Regulatory Impact Review and Final 

Environmental Impact Statement for 

Corals and Reef Associated Plants 

and Invertebrates of Puerto Rico 

and the U. S. Virgin Islands, 60 FR 

58221. Available on-line at: <www.

caribbeanfmc.com/coralFMP.htm>

Carpenter KE (ed) (2002) The living marine 

resources of the Western Central 

Atlantic. Volume 1: Introduction, 

molluscs, crustaceans, hagfishes, 

sharks, batoid fishes, and chimaeras. 

In FAO Species Identification Guide 

for Fishery Purposes and American 

Society of Ichthyologists and 

Herpetologists Special Publication 5. 

FAO Rome, pp 1375-2127

Chuenpagdee R, Morgan LE, Maxwell S, 

Norse EA, Pauly D (2003) Shifting 

gears: Assessing collateral impacts 

of fishing methods in the U.S. 

waters. Frontiers in Ecology and the 

Environment 1(10): 517–524

CITES (Convention on International Trade 

in Endangered Species of Wild 

Fauna and Flora) (2006). On-line 

species database of CITES-listed 

species. Accessed June 2006. 

Available on-line at: <http://sea.unepwcmc.org/isdb/CITES/Taxonomy/

?displaylanguage=eng>

CITES 2007. CITES trade database. On-line 

database. Accessed August 2007. 

Available on-line at: <http://www.unepwcmc.org/citestrade/trade.cfm>

Clark MR, Tittensor D, Rogers AD, Brewin 

P, Schlacher T, Rowden A, Stocks 

K, Consalvey M (2006). Seamounts, 

deep-sea corals and fisheries: 

vulnerability of deep-sea corals to 

fishing on seamounts beyond areas 

of national jurisdiction. UNEPWCMC, 

Cambridge, UK. 86 pp
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

334

Colin P (1978) Caribbean Reef Invertebrates 

and Plants. Neptune City, New Jersey: 

T.F.H. Publications, Inc.: 512pp

Correa T, Grasmueck M, Eberli G, Rathwell 

G, Viggiano D (2006) Morphology 

and Distribution of Deep-Water Coral 

Reefs in the Straits of Florida. Project 

prospectus: 3 pp

Cummings NJ, Matos-Caraballo D (2003) 

Summarized reported commercial 

landings in Puerto Rico from 1969-

2001 with specific notes on the Silk 

Snapper landings category. NOAA, 

NMFS, Caribbean Deepwater SEDAR 

Data Workshop Report 4 SEDAR 

DOC-5: 14 pp

Dawson JP (2002) Biogeography of 

azooxanthellate corals in the 

Caribbean and surrounding areas. 

Coral Reefs 21 (1): 27-40

Dulvy NK, Freckleton RP, Polunin NV 

(2004) Coral reef cascades and the 

indirect effects of predator removal 

by exploitation. Ecology Letters 7: (5): 

410–416

EarthRef.org (2007) Seamount Catalog. World 

wide web digital archive, accessed 

August 2007. Available at: <http://

earthref.org/cgi-bin/er.cgi?s=sc-s0-

main.cgi>

FAO (Food and Agriculture Organization of 

the United Nations) (1993) Marine 

fishery resources of the Antilles: 

Lesser Antilles, Puerto Rico and 

Hispaniola, Jamaica, Cuba. FAO 

Fisheries Technical Paper 326: 235 

pp

FAO (2000) Fishery Country Profile for the 

Republic of Trinidad and Tobago. FAO 

FID/CP/TRI Rev 2 May 2000. World 

wide web article, accessed April 2005. 

Available on-line at: <http://www.fao.

org/fi/fcp/en/TTO/profile.htm>

FAO (2003) Report of the second Workshop 

on the Management of Caribbean 

Spiny Lobster Fisheries in the 

WECAFC Area. FAO Fisheries Report 

715

Freiwald A, Fosså JH, Grehan A, Koslow T, 

Roberts JM (2004) Cold-water coral 

reefs. UNEP-WCMC, Cambridge, UK: 

84 pp

Fratantoni DM (2001) North Atlantic surface 

circulation during the 1990’s observed 

with satellite-tracked drifters. Journal 

of Geophysical Research 106: 22067-

22093

Genin A, Dayton PK, Lonsdale PF, Spiess 

FN (1986) Coral on seamount peaks 

provide evidence of current acceleration 

over deep-sea topography. Nature 

322: 59-61

Gordon AL (1967) Circulation of the Caribbean 

Sea. Journal of Geophysical Research 

72: 6207-6223

Gordon J (2004) Deep-sea fisheries of the 

Atlantic Ocean. Presented at Deepsea Fisheries: Ecology, Economics 

and Conservation (September 12-14, 

2004) Woods Hole Oceanographic 

Institution and New England Aquarium. 

World wide web article, accessed 

August 2005 Available on-line at: 

<http://www.whoi.edu/institutes/oli/

activities/symposia-deepsea-gordon.

pdf>

Gosner KL (1978) Peterson Field Guides, 

Atlantic Seashore from the Bay of 

Fundy to Cape Hatteras, Houghton 

Mifflin, 336 pp

Greenberg H, Greenberg D (2006) The 

U.S. Virgin Islands Alive (second 

edition). Hunter Travel Guides, Hunter 

Publishing Inc, Edison, NJ: 322 pp
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

335

Gunn JT, Watt DR (1982) On the currents 

and water masses north of the Antilles/

Bahamas Arc. Journal of Marine 

Research 40: 1-48

Gyory J, Mariano AJ, Ryan EH (2005a) “The 

Caribbean Current” Ocean Surface 

Currents. World wide web article, 

accessed March 2005 Available online at: <http://oceancurrents.rsmas.

miami.edu/caribbean/caribbean.html>

Gyory J, Rowe E, Mariano AJ, Ryan EH 

(2005b) “The Florida Current” Ocean 

Surface Currents. World wide web 

article, accessed March 2005 Available 

on-line at: <http://oceancurrents.

rsmas.miami.edu/caribbean/florida.

html>

Gyory J, Mariano AJ, Ryan EH (2005c) 

“The Loop Current” Ocean Surface 

Currents. World wide web article, 

accessed March 2005. Available online at: <http://oceancurrents.rsmas.

miami.edu/caribbean/loop-current.

html>

Gyory J, Mariano AJ, Ryan EH (2005d) “The 

Yucatan Current” Ocean Surface 

Currents. World wide web article, 

accessed March 2005. Available online at: <http://oceancurrents.rsmas.

miami.edu/caribbean/yucatan.html>

Harbison GR, Janssen J (1987) Encounters 

with a Swordfish (Xiphias gladius) and 

Sharptail Mola (Masturus lanceolatus) 

at Depths Greater Than 600 Meters. 

Copeia 1987 (2): 511-513

Hartman WD (1973) Beneath Caribbean 

Reefs. Discovery 9: 13-26

Haughton M, Mahon R, McConney P, Kong 

GA, Mills A (2004) Establishment of 

the Caribbean Regional Fisheries 

Mechanism. Marine Policy 28: 351-

359

Humann P (1993) Reef Coral Identification: 

Florida, Caribbean, Bahamas. 

Jacksonville, Florida: New World 

Publications, Inc.: 239 pp

James NP, Ginsburg RN (1979) The Seaward 

Margin of Belize Barrier and Atoll Reefs. 

Special Publication 3, International 

Association of Sedimentologists, 

Blackwell Science Publishing, Oxford, 

London, Edinburgh, Melbourne: 203 

pp

Johns WE, Townsend TL, Fratantoni DM, 

Wilson WD (2002) On the Atlantic 

inflow to the Caribbean Sea. DeepSea Res I (49): 211–243

Kapela W, Lasker HR (1999) Size dependent 

reproduction in the Caribbean 

gorgonian Pseudoplexaura porosa. 

Marine Biology 135:107-114

Keller NB, Pasternak FA, Naumov DV 

(1975) Bottom deep-sea Coelenterata 

from the Caribbean Sea and Gulf 

of Mexico (from material from the 

14th expedition of the `Akademik 

Kurchatov’). In: Scientific studies 

Caribbean Sea, Gulf of Mexico and 

adjacent waters. Transactions of the 

P.P. Shirshov Institute of Oceanology 

(Trudy Instituta okeanologii im. P.P. 

Shirshova) 100: 147-159 [in Russian 

with English summary]

Kojis B (2004) Census of the Marine 

Commercial Fishers of the U. S. Virgin 

Islands. U.S. Virgin Islands Division of 

Fish and Wildlife: 91 pp

Kojis B (2005) NOAA Fisheries - State Marine 

Fisheries Directors’ Biennial Meeting 

2005 - US Caribbean. 2005 State 

Directors’ meeting, April 11-13, 2005. 

PDF presentation
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

336

Koslow JA, Boehlert GW, Gordon JDM, 

Haedrich RL, Lorance P, Parin N 

(2000) Continental slope and deepsea fisheries: implications for a fragile 

ecosystem. ICES Journal of Marine 

Science 57 (3): 548-557

Lang JC (1974). Biological zonation at the 

base of a reef. Amer. Sci., 62 (3): 272- Amer. Sci., 62 (3): 272-

281

Lang JC, Hartman WD, Land LS (1975). 

Sclerosponges: Primary framework 

constructors on the Jamaican forereef. Journal of Marine Research 33 

(2): 952-959

Larsen K (1999) Deep-sea tanaidacean 

(Crustacea: Peracarida) from the 

Albatross cruises 1885-86, with keys 

to the suborder Neotanaidomorpha. 

Journal of Natural History 33 (8): 

1107-1132

Lee TN, Johns WE, Zantopp R, Fillenbaum 

ER (1996) Moored observations of 

western boundary current variability 

and thermohaline circulation 26.5°N in 

the subtropical North Atlantic, Journal 

of Physical Oceanography 26: 962-

963

Le Goff-Vitry MC (2003) Molecular ecology of 

the deep-sea coral Lophelia pertusa

in the North East Atlantic. PhD thesis. 

University of Southampton.

Littler MM, Littler DS (1999) The First 

Oceanographic Expedition to Navassa 

Island. Reef Encounter 25: 26-30

Melton International Tackle 2006 Master 

Catalog (2007). Fishing tackle and 

accessories catalog, 182 pp

Messing CG, Neumann AC, Lang JC (1990) 

Biozonation of deep water lithoherms 

and associated hardgrounds in the 

northeastern Straits of Florida. Palaios 

5: 15-33

Merrett NR, Haedrich RL (1997) DeepSea Demersal Fish and Fisheries. 

Chapman and Hall Great Britain: 296 

pp

Miller MW (2003) Status of reef resources 

of Navassa Island: Cruise report Nov. 

2002. NOAA Technical Memorandum 

NMFS-SEFSC-501: 119 pp

Morgan LE, Tsao CF, Guinotte JM (2006) 

Status of deep sea corals in U.S. 

waters, with recommendations for 

their conservation and management. 

Marine Conservation Biology Institute, 

Bellevue, Washington: 64 pp

Mullins HT, Newton CR, Heath K, Van Buren 

HM (1981) Modern deep-water coral 

mounds north of Little Bahama Bank; 

criteria for recognition of deep-water 

coral bioherms in the rock record. J 

Sed Res 51 (3): 999-1013

Mumby PJ, Dahlgren CP, Harborne AR, 

Kappel CV, Micheli F, Brumbaugh DR, 

Holmes KE, Mendes JM, Broad K, 

Sanchirico J, Buch K, Box S, Stoffle 

RW, Gill AB (2006) Fishing, Trophic 

Cascades, and the Process of Grazing 

on Coral Reefs. Science, Vol. 311 

(5757): 98 – 101

McNeill D (2006) Dimensional Aspects of 

Deepwater Mounds, eastern Straits of 

Florida. Project prospectus, 3 pp

Nelson R, Appeldoorn DS (1985) A 

submersible survey of the continental 

slope of Puerto Rico and the U.S. 

Virgin Islands, Cruise Report, RV 

Seward Johnson, October 1–23: 76 

pp

Neumann AC, Ball MM (1970) Submersible 

observations in the Straits of Florida: 

geology and bottom currents. 

Geological Society of America Bulletin 

81: 2861-2874
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

337

Neumann AC, Kofoed JW, Keller GH (1977) 

Lithoherms in the Straits of Florida. 

Geology 5 (1): 4-10

Nizinski M, Ross S (2002) Views Forward, 

Views Aft, August 14, 2002. On-line 

research cruise log dated August 14, 

2002. Accessed December, 2006, 

Available on-line at: <http://www.

oceanexplorer.noaa.gov/explorations/

02sab/logs/aug14/aug14.html>

NOAA (National Oceanic and Atmospheric 

Administration) (2005) Deep-Sea 

Coral Collection Protocols. NOAA 

Technical Memorandum. Deep-Sea 

Coral Panel of Experts. Draft dated 

January, 2005: 56 pp

NOAA (2007) Benthic Habitat Characterization 

and Bathymetry of Mid-water Habitats 

in the USVI and Puerto Rico (2007). 

Project on-line prospectus, accessed 

July 2007. Available on-line at: <http://

www8.nos.noaa.gov/nccos/npe/

projectdetail.aspx?id=402&fy=2007>

OBIS (Ocean Biogeographic Information 

System) (2006) On-line database, 

accessed July 2006. Available on-line 

at: <www.iobis.org> 

Offshore Fishing Forum (2006) The Marlin 

Club, Offshore Fishing Forum. Message 

posted by WahooKing on January 13, 

2005. Retrieved from the world wide 

web February 2006. Available on-line 

at: <http://www.marlinnut.com/cgi-bin/

dcforum/dcboard.cgi?az=printer_form

at&om=639&forum=DCForumID3 >

Opresko DM (1972) Redescriptions of 

antipatharians described by L.F. 

Pourtales. Bulletin of Marine Science 

22(4): 950-1017

Opresko DM (1974) A study of the classification 

of the Antipatharia (Coelenterata: 

Anthozoa), with redescriptions of 

eleven species. Ph.D. Dissertation, 

University of Miami: 149 pp

Opresko DM (1996) New species of black coral 

(Cnidaria: Anthozoa: Antipatharia) 

from the Caribbean, Bulletin of Marine 

Science 58: 289-300

Opresko DM (2002) Revision of the 

Antipatharia (Cnidaria: Anthozoa). 

Part II. Schizopathidae. Zoologische 

Mededelingen, Leiden. 76: 411–442

Opresko DM (2003) Revision of the 

Antipatharia (Cnidaria: Anthozoa). 

Part III. Cladopathidae. Zoologische 

Mededelingen, Leiden 77 (31): 495-

536

Opresko DM (2006) Antipatherian Corals 

Reported from the Caribbean. 

Unpublished list

Opresko DM, Cairns SD (1994) Description 

of the new genus Allopathes (Cnidaria: 

Antipatharia) and its type species 

Cirripathes desbonni. Proceedings of 

the Biological Society of Washington 

107(1): 185-192

Opresko DM, Sanchez JA (2005) Caribbean 

Shallow-water Black Corals (Cnidaria: 

Anthozoa: Antipatharia). Caribbean 

Journal of Science 41 (3): 492-507

Pasternak FA (1975) Deep-sea pennatularian 

genus Umbellula from the Caribbean 

Sea and Puerto-Rican Trench. In: 

Scientific studies Caribbean Sea, 

Gulf of Mexico and adjacent waters. 

Transactions of the P.P. Shirsov 

Institute of Oceanology 100: 160-173 

[in Russian with English summary]

Peters EC, Cairns SD, Pilson MEQ, Wells 

JW, Jaap WC, Lang JC, Vasleski 

CE, St. Pierre Gollahon L (1988) 

Nomenclature and biology of Astrangia 

poculata (Ellis and Solander, 1786). 

Proceedings of the Biological Society 

of Washington 101(2): 234-250
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

338

Picton BE, Howson CM (eds) (2000) The 

species directory of the marine fauna 

and flora of the British Isles and 

surrounding seas. Ulster Museum and 

The Marine Conservation Society, 

Belfast and Ross-on-Wye. CD-ROM 

Edition. On-line photographic guide, 

page: Funiculina quadrangularis

(Pallas, 1766). Accessed November 

2006. Available on-line at: <http://www.

seaslug.org.uk/marinelife/species.

asp?item=D10500>

Pillsbury JE (1890) The Gulf Stream - A 

description of the methods employed 

in the investigation, and the results of 

the research. USCC-Geodetic Survey 

Silver Spring MD, pp 461-620

Pourtalès LF de (1867) Contributions to the 

Fauna of the Gulf Stream at great 

depths. Bulletin of the Museum of 

Comparative Zoology, Harvard, 1: 

121-142

Pourtalès LF de (1871) Deep-sea corals. 

Illustrated catalogue of the Museum 

of Comparative Zoology at Harvard 

College 4: 93 pp

Pourtalès LF de (1874) Zoological results 

of the Hassler Expedition, deep-sea 

corals Illustrated Catalogue of the 

Museum of Comparative Zoology 8: 

33-49

Pourtalès LF de (1880) Report on the results 

of dredging, under the supervision of 

Alexander Agassiz, in the Caribbean 

Sea, 1878-79, by the United States 

Coast Survey steamer “Blake”. VI. 

Report on the corals and Antipatharia. 

Bulletin of the Museum of Comparative 

Zoology 6 (4): 95-120

Prado J, Drew S (1999) Research and 

Development in Fishing Technology in 

Latin America. FAO Fisheries Circular 

944 FIIT/x2173

Reed JK (2002) Comparison of deepwater coral reefs and lithoherms off 

southeastern U.S.A. Hydrobiologia 

471: 43-55

Reed JK, Shepard A, Koenig C, Scanlon K, 

Gilmore G (2005). Mapping, habitat 

characterization, and fish surveys 

of the deep-water Oculina coral reef 

Marine Protected Area: a review of 

historical and current research. In: 

Freiwald A, Roberts J (Eds), Cold-water 

Corals and Ecosystems. Proceedings 

of the 2nd International Symposium 

on Deep-Sea Corals Sept. 9-12 2003 

Erlanger Germany. Springer-Verlag 

Berlin Heidelberg, 1243 pp

Reed JK (2006) Final Cruise Report, 

Bahamas-Florida Deep-Water Reef 

Expedition: Discovery and Exploration 

of Deep-Water Reefs in the Straits of 

Florida, May 22-30, 2006. Submitted 

to: The Bahamas Department of 

Fisheries- Ministry of Agriculture, 

Fisheries, and Local Government, 

DVD (Cruise Report and images) 

Reyes J, Santodomingo N, Gracia A, 

Borrero-Pérez G, Navas G, Ladino 

LM, Bermúdez A, Benavides M (2005) 

Southern Caribbean azooxanthellate 

coral communities off Colombia. In: 

Freiwald A, Roberts J (Eds) Cold-water 

Corals and Ecosystems Proceedings 

of the 2nd International Symposium 

on Deep-Sea Corals Sept. 9-12 2003 

Erlanger Germany. Springer-Verlag Springer-Verlag 

Berlin Heidelberg, 1243 pp

Rezek R, Bright TJ, McGrail DW (1985) Reefs 

and banks of the northwestern Gulf of 

Mexico. Their geological, biological, 

and physical dynamics. John Wiley 

and Sons, New York: 259 pp

Roberts CM (2002) Deep impact: the rising 

toll of fishing in the deep sea. Trends 

in Ecology & Evolution 17: 242-245
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

339

Rogers AD (1999) The biology of Lophelia 

pertusa and other deep-water reefforming corals and impacts from 

human activities. International Review 

of Hydrobiology 84 (4): 315-406

Rowe E, Mariano AJ, Ryan EH (2005) “The 

Antilles Current” Ocean Surface 

Currents. World wide web article, 

accessed April 2005. Available online at: <http://oceancurrents.rsmas.

miami.edu/atlantic/antilles.html>

Sala E, Ballesteros E, Starr RM (2001) Rapid 

decline of Nassau grouper spawning 

aggregations in Belize: fishery 

management and conservation needs. 

Fisheries 26: 23-30

Santodomingo N, Reyes J, Gracia A, Martínez 

A, Ojeda G, García G (2006) Madracis 

coral communities off San Bernardo 

Islands (Colombian Caribbean). 

Bulletin of Marine Science, in press, 

manuscript received November 2006, 

29 pp

Sedberry GR, Ulrich GF, Applegate AJ (1994) 

Development and status of the fishery 

for wreckfish (Polyprion americanus) 

in the southeastern United States. 

Proceedings of the Gulf and Caribbean 

Fishery Institute 43: 168-192

Sheridan P, Hill R, Kojis B (2006) Trap Fishing 

in the U.S. Virgin Islands: How and 

Where Effort is Exerted. Proceedings 

of the Gulf and Caribbean Fisheries 

Institute 57: 175-188

South Atlantic Fishery Management Council 

(1998) Final Habitat Plan for the 

South Atlantic Region: Essential 

Fish Habitat Requirements for 

Fishery Management Plans of the 

South Atlantic Fishery Management 

Council. 457 pp. Available on-line 

at: <http://www.safmc.net/Default.

aspx?tabid=80>

Spalding MD (2004) A guide to the Coral 

reefs of the Caribbean. University of 

California Press: 256 pp

Squires DF (1964) Fossil coral thickets in 

Wairarapa, New Zealand: Journal of 

Paleontology 30 (5): 904-915

Staiger JC (1968a) Narrative of the Cruise p6802 to the Northwestern Caribbean, 

March 11-26, 1968. Processed Report 

Marine Invertebrate Museum RSMAS 

University of Miami: 24 pp

Staiger JC (1968b) Narrative of the Cruise p6806 to the Southern Caribbean, July 

6 - August 3, 1968. Processed Report 

Marine Invertebrate Museum RSMAS 

University of Miami: 71 pp

Staiger JC (1969) Narrative of the Cruise p6806 - Antillean Arc, June 27 - July 

31, 1969, 1968. Processed Report, 

RSMAS, University of Miami, 65 pp

Staiger JC (1971) Narrative of the Cruise p7101 - Central America, January 20 

- 5 February 1971. Processed Report 

Marine Invertebrate Museum RSMAS 

University of Miami: 31pp

Staiger JC, Voss GL (1970) Narrative of 

the Cruise p-7006 - Hispaniola and 

Jamaica, June 26 - July 23, 1970. 

Processed Report Marine Invertebrate 

Museum RSMAS University of Miami: 

57 pp

Steneck RS, Sala E (2005). Large Marine 

Carnivores: Trophic Cascades and TopDown Controls in Coastal Ecosystems 

Past and Present. In: Large Carnivores 

and the Conservation of Biodiversity, 

Ray JC, Redford KH, Steneck RS, 

Berger J (Eds). Island Press: 512 pp

Stetson TR, Squires DF, Pratt RM (1962) 

Coral Banks Occurring in Deep Water 

on the Blake Plateau. American 

Museum Novitates 2114: 1-39
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

340

Sword Fishing Central (2006) SFC Fishing 

Forums Deep Dropping sub forum. 

Deep drop weights? Discussion 

thread. Retrieved from the world wide 

web November 2006. Available on-line 

at: <http://www.swordfishingcentral.

com/forum/deep-dropping/6194-

deep-drop-weights.html>

Swingle WE, Dammann AE, Yntema JA 

(1970) Survey of the Commercial 

Fishery of the Virgin Islands of the 

United States. Proceedings of the 

Gulf and Caribbean Fisheries Institute 

22:110-121.

Theile S (2001) Queen Conch fisheries and 

their management in the Caribbean. 

Technical Report to the CITES 

Secretariat. Brussels, TRAFFIC 

Europe, 95 pp

Tobias W (2004) Status of the Reefs: Reef 

Fish Resources. U.S. Virgin Islands 

Department of Planning and Natural 

Resources, Division of Fish and 

Wildlife: 11 pp

UNEP-WCMC (United Nations Environment 

Programme-World Conservation 

Monitoring Centre) (2003) Checklist 

of fish and invertebrates listed in 

the CITES appendices and in EC 

Regulation 338/97. 6th Edition. JNCC 

Report 341

van der Land J, Opresko DM (2001) 

Antipatharia, in: Costello MJ et al. 

(eds.) (2001). European register 

of marine species: a check-list of 

the marine species in Europe and 

a bibliography of guides to their 

identification. Collection Patrimoines 

Naturels 50: 109 pp

Voss GL (1966a) Narrative of the Cruise p6607 to the Southwestern Caribbean, 

July 7 - 22. Processed Report, RSMAS, 

University of Miami: 38 pp

Voss GL (1966b) Biological Survey the 

Southwestern Caribbean. Processed 

Report, RSMAS, University of Miami: 

36 pp

Voss GL (1971) Narrative of the Cruise p7106 to the Nares Abyssal Basin and 

Puerto Rico Trench, 26 June - 13 July 

1971. Processed Report, RSMAS, 

University of Miami: 32 pp

Veronique PT (1987) Annotated Checklist 

of the Gorgonacea from Martinique 

and Guadeloupe Islands (F.W.I.). Atoll 

Research Bulletin 303: l-16.

Viada ST, Cairns SD (1987) Range extensions 

of ahermatypic Scleractinia in the Gulf 

of Mexico. Northeast Gulf Science 9 

(2): 131-134

Watling L, Auster PJ (2005) Distribution 

of deep-water Alcyonacea off the 

Northeast coast of the United States. 

In: Freiwald A, Roberts JM (eds) 

Cold-Water Corals and Ecosystems, 

Springer-Verlag: 259-276

Warner GF (1981) Species descriptions and 

ecological observations of black corals 

(Antipatharia) from Trinidad. Bulletin 

of Marine Science 31: 147-163

Wilber JR, Neumann AC (1993) Effects of 

submarine cementation on microfabrics 

and physical properties of carbonate 

slope deposits, Northern Bahamas. In: 

Rezak R, Lavoie DL (eds). Carbonate 

Microfabrics, Springer, New York: 79-

94

Wilkinson C (ed) (2004) Status of coral 

reefs of the world: 2004. Volume 2. 

Australian Institute of Marine Science, 

Townsville, Queensland, Australia: 

301 pp
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Appendix 8.1. Deep-sea corals of the wider Caribbean region, including those located in the U.S. Caribbean▲ Bold denotes major and potential structure-forming scleractinia (species associated with deep water banks, lithoherms, etc.) and potential habitat-forming antipatharia, octocorallia, pennatulacea, and hydrozoa (species greater than or equal to 25 cm species greater than or equal to 25 cm in height or width ♠ Bold denotes distribution including U.S. Caribbean territories‡ For material examined: HBOI = Harbor Branch Oceanographic Institute; USNM = United States National Museum

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Phylum CnidariaClass AnthozoaSubclass Hexacorallia (Zoantharia)Order Scleractinia

Caryophyllidae Anomocora fecunda

Widespread Caribbean and Bahamas, including the U.S. Virgin Islands 37-640

Cairns 1979, 2000, UNEP-WCMC 2003; Reyes et al. 2005

Caryophyllidae Anomocora marchadi Straits of Florida, southern Caribbean 35-229

Cairns 1979, 2000; Cairns and Zibrowis 1997

Caryophyllidae Anomocora prolifera

Straits of Florida, southern Caribbean, Lesser Antilles 30-329

Cairns 1979, 2000; Reyes et al. 2005

Anthemiphylliidae Anthemiphyllia patera

Straits of Florida, Bahamas, Lesser Antilles 500-700 Cairns 1979

Rhizangiidae Astrangia poculata

Throughout Caribbean, including Puerto Rico 0-263 Cairns 1999, 2000

Dendrophylliidae Balanophyllia bayeri Mexico, Colombia 274-311 Cairns 1979; Reyes et al. 2005Dendrophylliidae Balanophyllia cyathoides

Straits of Florida, Yucatan Channel, Lesser Antilles; Venezuela 45-494

Cairns 1979, 2000; Reyes et al. 2005

Dendrophylliidae Balanophyllia dineta

Southeastern Caribbean, Lesser Antilles 27-274 Cairns 2000

Dendrophylliidae Balanophyllia floridana

Straits of Florida, southeastern and western Caribbean 13-220 Cairns 2000

Dendrophylliidae Balanophyllia hadros Only off Nicaragua 238-274 Cairns 1979Dendrophylliidae Balanophyllia palifera

Bahamas, Antilles, Yucatan Channel, Colombia 53-708

Cairns 1979, 2000; Reyes et al. 2005
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Dendrophylliidae Balanophyllia wellsi

Bahamas, Cuba, Grenada, Jamaica, Colombia 412-505

Cairns 1979, UNEP-WCMC 2003; Reyes et al. 2005

Dendrophylliidae Bathypsammia fallosocialis Straits of Florida, St. Lucia 244-805 Cairns 1979Caryophyllidae Caryophyllia ambrosia caribbeana

Widespread Caribbean, Bahamas, including U.S. territories 183-1646 Cairns 1979; Reyes et al. 2005

Caryophyllidae Caryophyllia antillarum

Straits of Florida, Bahamas, Antilles, including US territories 150-730 Cairns 1979, 2000

Caryophyllidae Caryophyllia barbadendis Barbados; Colombia 129-249

Cairns 1979, 2000; Reyes et al. 2005

Caryophyllidae Caryophyllia berteriana

Widespread Caribbean, Bahamas, including U.S. territories 99-1033

Cairns 1979, 2000; Reyes et al. 2005

Caryophyllidae Caryophyllia corrugata

Antilles, Virgin Islands to Cuba, including U.S. territories 183-380 Cairns 1979

Caryophyllidae Caryophyllia crypta

Bahamas and Caribbean, Greater Antilles, southern and western Caribbean, including U.S. territories 12-183 Cairns 2000

Caryophyllidae Caryophyllia parvula Antilles, Venezuela 97-399 Cairns 1979Caryophyllidae Caryophyllia paucipalata

Lesser Antilles, including U.S. territories 714-843 Cairns 1979

Caryophyllidae Caryophyllia polygona Straits of Florida, Antilles 700-1817 Cairns 1979Caryophyllidae Caryophyllia zopyros Antilles 73-618 Cairns 1979, 2000Caryophyllidae Cladocora debilis

Straits of Florida, eastern and southern Caribbean, including Puerto Rico

32-480 (common 

50-100)

Cairns 2000; UNEP-WCMC 2003; Reyes et al. 2005

Dendrophylliidae Cladopsammia manuelensis Straits of Florida, St. Lucia 70-366 Cairns 1979, 2000Caryophyllidae Coenocyathus caribbeana Bahamas and Caribbean 5-100 Cairns 2000
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Caryophyllidae Coenocyathus parvulus

Bahamas and Caribbean, including U.S. territories 97-399

Cairns 1979, 2000; Reyes et al. 2005

Caryophyllidae Coenosmilia arbuscula

Widespread Caribbean, Bahamas, including U.S. territories 74-622

Cairns 1979, 2000; Reyes et al. 2005

Caryophyllidae Colangia immersa

Throughout Caribbean, Bahamas, including U.S. territories 0.5-347 Cairns 2000

Caryophyllidae Concentrotheca laevigata Straits of Florida, Yucatan Channel 183-800 Cairns 1979Turbinoliidae Cylicia inflata Only off Havana, Cuba 183-443 Cairns 1979Caryophyllidae Dasmosmilia lymani

Straits of Florida, Cuba, Grenada, Venezuela 37-366

Cairns 1979, 2000, UNEP-WCMC 2003

Caryophyllidae Dasmosmilia variegata

Straits of Florida, southern Lesser Antilles 110-421 Cairns 1979, 2000

Turbinoliidae Deltocyathoides stimpsonii Straits of Florida, Lesser Antilles 110-553 Cairns 1979, 2000Caryophyllidae Deltocyathus agassizii Straits of Florida, Anguilla 494-907 Cairns 1979Caryophyllidae Deltocyathus calcar

Widespread Caribbean, Bahamas, including U.S. territories 81-675

Cairns 1979, 2000; Reyes et al. 2005

Caryophyllidae Deltocyathus eccentricus

Widespread Caribbean, Bahamas, including U.S. territories 183-907 Cairns 1979

Caryophyllidae Deltocyathus italicus

Widespread Caribbean, Bahamas, including U.S. territories 403-2634 Cairns 1979

Caryophyllidae Deltocyathus moseleyi

Straits of Florida, Belize, Lesser Antilles 201-777 Cairns 1979

Caryophyllidae Deltocyathus pourtalesi Straits of Florida, Bahamas 311-567 Cairns 1979Dendrophylliaidae Dendrophyllia alternata Antilles, Bahamas 276-900 Cairns 1979Caryophyllidae

Desmophyllum dianthus (=D. cristagalli)

Straits of Florida, Bahamas, Martinique, U.S. Virgin Islands 155-2200

Cairns 1979, 2000, UNEP-WCMC 2003

Caryophyllidae Desmophyllum striatum

Greater Antilles, Bahamas, including U.S. territories 277-823 Cairns 1979, 2000

Dendrophylliaidae Eguchipsammia cornucopia

Bahamas, Straits of Florida, Yucatan Channel, southeastern Caribbean 93-300

Cairns 1979, 2000; Reyes et al. 2005

Dendrophylliaidae Eguchipsammia gaditana Yucatan Channel, Venezuela 97-505 Cairns 1979, 2000
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Dendrophylliaidae Enallopsammia profunda Straits of Florida, St. Lucia 403-1748 Cairns 1979Dendrophylliaidae Enallopsammia rostrata

Straits of Florida, Bahamas, Antilles, western Caribbean, Antilles, including U.S. Virgin Islands 300-1646 Cairns 1979, UNEP-WCMC 2003

Flabellidae Flabellum (pavoninum) atlanticum Bahamas, Cuba 357-618 Cairns 1979, UNEP-WCMC 2003Flabellidae Flabellum floridanum

Western Straits of Florida, Mexico, Panama 80-366

Cairns 1979, 2000; UNEP-WCMC 2003

Flabellidae Flabellum moseleyi

Widespread Caribbean, including U.S. territories 216-1097 Cairns 1979

Fungiacyathidae Fungiacyathus marenzelleri Bahamian archipelago 1450-2745 Cairns 1979Fungiacyathidae Fungiacyathus pusillus

Straits of Florida, Yucatan, Lesser Antilles 285-439 Cairns 1979

Fungiacyathidae Fungiacyathus symmetricus Widespread Caribbean, Bahamas. 183-1664 Cairns 1979Gardineriidae Gardineria minor Throughout Caribbean, Bahamas 2-146 Cairns 1979, 2000Gardineriidae Gardineria paradoxa

Antilles, Yucatan Channel, Straits of Florida 97-700 Cairns 1979, 2000

Gardineriidae Gardineria simplex Bahamas, Antilles, Yucatan Channel 46-241 Cairns 2000Guyniidae Guynia annulata

Bahamas, Straits of Florida, western Caribbean, Antilles 30-653 Cairns 1979, 2000

Flabellidae Javania cailleti 

Widespread Caribbean, Bahamas, including U.S. territories 30-1809

Cairns 1979, 2000; Reyes et al. 2005

Flabellidae Javania pseudoalabastra Bahamas, Jamaica 1089-1234 Cairns 1979Caryophyllidae Labyrinthocyathus langae

Straits of Florida, Bahamas, Antilles, western Caribbean 695-810 Cairns 1979

Micrabaciidae Leptopenus discus Only off south east Cuba 2842-3475 Cairns 1979Caryophyllidae Lophelia pertusa

Straits of Florida, Bahamas, southern Caribbean, Antilles, including Puerto Rico and U.S. Virgin Islands 146-1200

Cairns 1979, 2000, UNEP-WCMC 2003

Pocilloporidae Madracis asperula

Widespread Caribbean, including U.S. territories (absent Bahamas) 24-311 Cairns 2000; Reyes et al. 2005

Pocilloporidae Madracis brueggemanni Southern Caribbean 51-130 Cairns 2000
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Pocilloporidae Madracis myriaster

Widespread Caribbean, Bahamas, including Puerto Rico and U.S. Virgin Islands 20-1220

Cairns 1979, 2000, UNEP-WCMC 2003

Pocilloporidae

Madracis pharensis formapharensis

Widespread Caribbean, Bahamas, including U.S. territories 11-333 Cairns 2000; Reyes et al. 2005

Oculinidae Madrepora carolina

Greater Antilles, Bahamas, Tobago, western Caribbean, including Puerto Rico and U.S. Virgin Islands, Venezuela

53-801 (common 

100-300)

Cairns 1979, 2000; UNEP-WCMC 2003; Reyes et al. 2005

Oculinidae Madrepora oculata

Widespread Caribbean, Bahamas, including Puerto Rico and U.S. Virgin Islands 144-1391 Cairns 1979

Oculinidae Oculina tenella Straits of Florida, Cuba

25-159 (common 

40-60) Cairns 2000; UNEP-WCMC 2003

Caryophyllidae Oxysmilia rotundifolia

Widespread Caribbean, Bahamas, including U.S. territories 46-640

Cairns 1979, 2000; Reyes et al. 2005

Caryophyllidae Paracyathus pulchellus

Widespread Caribbean, including U.S. territories 17-250

Cairns 1979, 2000; Reyes et al. 2005

Turbinoliidae Peponocyathus folloculus Straits of Florida, Antilles 284-457 Cairns 1979Caryophyllidae Phacelocyathus flos Bahamas, Antilles, western Caribbean 20-355 Cairns 1979, 2000Flabellidae Placotrochides frusta Lesser Antilles 497-907 Cairns 1979Caryophyllidae Polycyathus mayae

Straits of Florida, Bahamas, northern Caribbean, including U.S. territories, Barbados 137-309 Cairns 2000; Reyes et al. 2005

Caryophyllidae Polycyathus senegalensis

Disjunct distribution, Florida, southern Caribbean 12-143 Cairns 2000

Flabellidae Polymyces fragilis

Straits of Florida, Bahamas, western and southern Caribbean, southern Lesser Antilles 75-822

Cairns 1979, 2000; Reyes et al. 2005

Guyniidae Pourtalocyathus hispidus

Straits of Florida, Bahamas, Antilles, including U.S. territories 349-1200 Cairns 1979

Caryophyllidae Premocyathus cornuformis

Straits of Florida, Bahamas, northern and eastern Caribbean 137-931 Cairns 1979, 2000
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Dendrophylliaidae Rhizopsammia goesi

Lesser Antilles, Colombia, Bahamas, including U.S. Virgin Islands 4.5-119

Cairns 1977, 2000, UNEP-WCMC 2003

Caryophyllidae Rhizosmilia gerdae

Straits of Florida, Bahamas, Yucatan Channel, northeastern Antilles, including U.S. territories 123-549 Cairns 1979, 2000

Caryophyllidae Rhizosmilia maculata

Widespread Caribbean, Bahamas, including U.S. territories 0.5-508 Cairns 2000

Guyniidae Schizocyathus fissilis

Straits of Florida, Bahamas, western Caribbean, Antilles (from Puerto Ricoto Grenada) 88-640 Cairns 1979, 2000

Caryophyllidae Solenosmilia variabilis

Straits of Florida, Lesser Antilles, Jamaica, southern Caribbean 220-1383 Cairns 1979

Guyniidae Stenocyathus vermiformis

Straits of Florida, western Caribbean, Bahamas, Antilles, including U.S. territories 165-835 Cairns 1979, 2000

Caryophyllidae Stephanocyathus coronatus

Throughout Caribbean, Bahamas, including U.S. territories 543-1250 Cairns 1979

Caryophyllidae Stephanocyathus diadema

Widespread Caribbean, including U.S. territories 795-2113 Cairns 1979

Caryophyllidae Stephanocyathus laevifundus

Antilles, Panama, Straits of Florida, and off Navassa Island 300-1158 Cairns 1979

Caryophyllidae Stephanocyathus paliferus Widespread Caribbean, Bahamas 299-715 Cairns 1979Caryophyllidae Tethocyathus cylindraceus

Straits of Florida, Jamaica, Bahamas, Barbados. 155-649 Cairns 1979

Caryophyllidae Tethocyathus recurvatus Yucatan Channel 320-488 Cairns 1979Caryophyllidae Tethocyathus variabilis

Straits of Florida, Yucatan, Jamaica, including U.S. territories 320-488 Cairns 1979

Caryophyllidae Thalamophyllia gombergi Only from Pourtales Terrace, Florida 155-220 Cairns 1979Caryophyllidae Thalamophyllia riisei

Bahamas, Antilles, including U.S. territories, Colombia 4-914

Cairns 1979, 2000; Reyes et al. 2005

Dendrophylliidae Thecopsammia socialis Gerogia to FL Keys, Bahamas 214-878 Cairns 1979; UNEP-WCMC 2003Turbinoliidae Trematotrochus corbicula Only northwest Cuba 400-576 Cairns 1979
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Caryophyllidae Trochocyathus fossulus

Bahamas, Virgin Islands, including U.S. territories 205-380 Cairns 1979

Caryophyllidae Trochocyathus rawsonii

Widespread Caribbean, Bahamas, including U.S. territories 55-700 Cairns 1979, 2000

Dendrophylliaidae Trochopsammia infundibulum northwestern Cuba, Lesser Antilles 532-1372 Cairns 1979

Order Zoanthidea

Gerardiidae Gerardia sp. Bahamas (Straits of Florida) >100 (~580-630) Messing et al. 1990

Order Antipatharia

Aphanipathidae Acanthopathes humilis

Bahamas, Barbados, Cuba, Grenada, Mexico, Montserrat, Saint Vincent and the Grenadines 129 - 491

Opresko 1972; UNEP-WCMC 2003; Opresko 2006

Aphanipathidae Acanthopathes thyoides

Cuba, Saint Vincent and the Grenadines ~40-240

Opresko 1972; UNEP-WCMC 2003; Opresko 2006

Antipathidae Allopathes desbonni

Barbados, Cuba, Guadeloupe, Montserrat 129-161

Opresko and Cairns 1994; UNEPWCMC 2003; Opresko 2006

Antipathidae Antipathes americana?

U.S. Virgin Islands, Grenada, Netherlands Antilles 37-532

Brook 1889; Opresko 1972; Warner 1981; OBIS 2006

Antipathidae Antipathes atlantica

Colombia, Jamaica, Mexico, Trinidad and Tobago 10 - 115

Warner 1981; UNEP-WCMC 2003; Reyes et al. 2005; Opresko 2006

Antipathidae Antipathes caribbeana

Bahamas, Colombia, Puerto Rico, Greater and Lesser Antilles

11->100 (common 

30-60)

Opresko 1996; UNEP-WCMC 2003; Opresko and Sanchez 2005

Antipathidae Antipathes columnaris

Greater and Lesser Antilles, includingU.S. Virgin Islands, Venezuela 73 - 567

Opresko 1974; UNEP-WCMC 2003; Reyes et al. 2005; Opresko 2006

Antipathidae Antipathes lenta

Straits of Florida, Cuba, Saint Vincent and the Grenadines, Trinidad and Tobago, Barbados, Honduras, Panama, Colombia, Venezuela 50-200

Opresko 1972; UNEP-WCMC 2003; Opresko and Sanchez 2005; Reyes et al. 2005; OBIS 2006; Opresko 2006
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Antipathidae Antipathes gracilis

from Colombia north throughout the Caribbean, including Panama, Trinidad and Tobago, and Colombia 20-160

Brook 1889, Warner 1981, STRI 2006; Reyes et al. 2005

Antipathidae Antipathes rhipidion? U.S. Virgin Islands Unk UNEP-WCMC 2003Antipathidae Antipathes tristis?

Barbados, Guadeloupe, Martinique, Montserrat, Saint Lucia Unk UNEP-WCMC 2003

Aphanipathidae Aphanipathes pedata Widespread Caribbean ~60-310 Opresko 1974; Opresko 2006Aphanipathidae Aphanipathes salix Guadeloupe, Colombia 107-333

Opresko 1972; UNEP-WCMC 2003; Reyes et al. 2005; Opresko 2006

Schizopathidae Bathypathes alternata

Straits of Florida (Bahamas), Yucatan Channel ~100-5000

Brook 1889 ; Opresko 1974; Opresko 2002; Opresko 2006

Schizopathidae Bathypathes patula

Mexico, Puerto Rico, Saint Kitts and Nevis, Caribbean Sea Basin, Cayman Trench, Yucatan Basin 100-5000

Brook 1889; Opresko 1974; Keller, et al. 1975; Opresko 2002; UNEP-WCMC 2003; OBIS 2006; Opresko 2006

Cladopathidae Chrysopathes sp. Widespread Caribbean >100 Opresko 2006Antipathidae Cirrhipathes spiralis?

Barbados, Cuba, Martinique, Montserrat, St. Vincent and the Grenadines UNEP-WCMC 2003

Aphanipathidae Distichopathes disticha Martinique, Mexico 95-190

Opresko 2004; CITES 2006; Opresko 2006

Aphanipathidae Distichopathes filix

Bahamas, Barbados, Cuba, Dominica, Guadeloupe, Martinique, Montserrat, SaintVincent and the Grenadines ~175-450

Opresko 2006; Opresko 1972; UNEP-WCMC 2003

Aphanipathidae Elatopathes abietina

Bahamas, Barbados, Cuba, Martinique, Mexico, Nicaragua, Saint Vincent and the Grenadines; Colombia 31 - 310

Opresko 1972; Rezak et al. 1985; UNEP-WCMC 2003; Reyes et al. 2005; Opresko 2006

Cladopathidae Heliopathes americana Widespread Caribbean >100 Opresko 2003; Opresko 2006Leiopathidae Leiopathes glaberrima

Straits of Florida, Jamaica, Mexico; Venezuela, Bahamas 176 - 549

Opresko 1974; UNEP-WCMC 2003; Opresko 2006; Material examined USNM 1026305
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Schizopathidae Parantipathes tetrasticha

Cuba, Guyana, Mexico, Puerto Rico, Saint Lucia 175 - 428

Opresko 1972; Opresko 2002; UNEP-WCMC 2003; Opresko 2006

Schizopathidae Phanopathes rigida

Bahamas, Barbados, Colombia, Cuba, Guadeloupe, Venezuela 64 - 640

Opresko 1972; Opresko 2002; UNEP-WCMC 2003; Opresko 2006

Myriopathidae Plumapathes pennacea

Throughout Caribbean, including U.S. Virgin Islands & Puerto Rico 3-229

Opresko 1974; Colin 1978; UNEP-WCMC 2003, CITES 2006; Opresko 2006

Antipathidae Stichopathes lutkeni

Barbados, Colombia, Dominican Republic, Trinidad and Tobago ~14-115

Brook 1889; Humann 1993; UNEP-WCMC 2003; Reyes et al. 2005; Opresko 2006

Antipathidae Stichopathes occidentalis Caribbean region ~20-160

Brook 1889; Opresko 2006; Reyes et al. 2005

Antipathidae Stichopathes pourtalesi

Barbados, Colombia, Cuba, Grenada, Martinique, Montserrat, Saint Vincent and the Grenadines >100

Pourtalès 1874, 1880; Brook 1889; Reyes et al. 2005; CITES 2006; Opresko 2006

Myriopathidae Tanacetipathes barbadensis Barbados, Trinidad and Tobago ~60-345

Brook 1889; van der Land and Opresko 2001; Warner 1981; UNEP-WCMC 2003; Opresko 2006

Myriopathidae Tanacetipathes hirta

Throughout Caribbean, including Puerto Rico 13 - 357

Opresko 1972; Colin 1978; Warner 1981; UNEP-WCMC 2003; Opresko 2006

Myriopathidae Tanacetipathes tanacetum

Throughout Caribbean, including Puerto Rico 46 - 915

UNEP-WCMC 2003; OBIS 2006; Opresko 2006

Myriopathidae Tanacetipathes thamnea Trinidad and Tobago ~45-345

Opresko 1970; Warner 1981; UNEP-WCMC 2003; Opresko 2006

Subclass OctocoralliaOrder Alcyonacea

Alcyoniidae Alcyonium digitatum Western Atlantic >200 Cairns 2005Alcyoniidae Anthomastus agassizi Straits of Florida, Bahamas 320-3186 Cairns 2005
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Alcyoniidae Anthomastus robusta var. delta Western Atlantic >200 Cairns 2005Alcyoniidae Bellonella rubistella Throughout Caribbean 24-329 Cairns 2005Clavulariidae Carijora (=Telesto?) operculata

Straits of Florida (off Havana, Cuba), probably throughout Antilles 60-298 Cairns 2005

Nephtheidae Drifta glomerata Western Atlantic >200 Cairns 2005Nephtheidae Duva florida Western Atlantic >200 Cairns 2005Nephtheidae Gersemia rubiformia Western Atlantic >200 Cairns 2005Nephtheidae Neospongodes portoricensis Bahamas, Caribbean 37-503 Humann 1993Nidaliiae Nidalia deichmannae Western Atlantic >200 Cairns 2005Nidaliiae Nidalia dissidens Western Atlantic >200 Cairns 2005; Reyes et al. 2005Nidaliiae Nidalia occidentalis Throughout Caribbean 37-311 Cairns 2005; Reyes et al. 2005Nidaliiae Nidalia rubrapunctata Western Atlantic ~82->200 Cairns 2005Nephtheidae Pseudodrifa capnella nigra

Straits of Florida (off Havana, Cuba), Bahamas 60-878

South Atlantic Fishery Management Council 1998; Cairns 2005; Reed 2006

Clavulariidae Scleranthelia rugosa rugosa Throughout Caribbean 175-586 Cairns 2005Nephtheidae Stereonepthya portoricensis Western Atlantic >200 Cairns 2005Clavulariidae Stereotelesto (=Telesto) corallina

Bahamas, Antilles, including Puerto Rico 23-188 Bayer 1961; Humann 1993

Clavulariidae Telesto nelleae

Straits of Florida (off Havana, Cuba), Bahamas 27-1153

Bayer 1961; South Atlantic Fishery Management Council 1998; Cairns 2005

Clavulariidae Telesto septentrionalis Western Atlantic >200 Cairns 2005Nephtheidae Trachythela rudis Western Atlantic 805 Cairns 2005

Order Gorgonacea

Isididae Acanella arbuscula Western Atlantic >200 Cairns 2005Isididae Acanella eburnea Western Atlantic >200 Cairns 2005Plexauridae Acanthactis austera Western Atlantic >200 Cairns 2005
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Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Plexauridae Acanthactis scabra Western Atlantic >200 Cairns 2005Acanthogorgiidae Acanthogorgia armata Western Atlantic >200 Cairns 2005Acanthogorgiidae Acanthogorgia aspera

Throughout Caribbean, including Puerto Rico >183

Watling and Auster 2005; Bayer 1961

Acanthogorgiidae Acanthogorgia schrammi

Straits of Florida; Guadeloupe; Colombia >200

Veronique 1987; Watling and Auster 2005; Reyes et al. 2005

Primnoidae Acanthoprimnoa goesi

Straits of Florida, Bahamas, Puerto Rico, Virgin Islands, including U.S. Virgin Islands 137-595

Cairns and Bayer 2004b; Cairns 2005

Primnoidae Acanthoprimnoa pectinata

northeastern Yucatan, lesser Antilles, Straits of Florida 194-686

Cairns and Bayer 2004b; Cairns 2005

Anthothelidae Anthothela bathybius Western Atlantic >200 Cairns 2005Anthothelidae Anthothela grandiflora Western Atlantic >200 Cairns 2005Anthothelidae Anthothela grandiflora sensu Western Atlantic >200 Cairns 2005Plexauridae Bebryce cinerea Western Atlantic >200 Cairns 2005Plexauridae Bebryce grandis Western Atlantic >200 Cairns 2005Plexauridae Bebryce parastellata Western Atlantic >200 Cairns 2005Plexauridae Caliacis nutans Western Atlantic >200 Cairns 2005Primnoidae

Callogorgia americana americana

Straits of Florida, Lesser Antilles, including U.S. territories 183-732

Cairns & Bayer 2002; Cairns 2005

Primnoidae Callogorgia gracilis

Throughout Caribbean, Bahamas, Antilles 82-514

Cairns & Bayer 2002; Cairns 2005

Primnoidae Callogorgia linguimaris Only Bahamas 1116

Cairns & Bayer 2002; Cairns 2005

Primnoidae Calyptrophora antilla Windward Passage 1399 Bayer 2001; Cairns 2005Primnoidae Calyptrophora gerdae Straits of Florida 229-556 Bayer 2001; Cairns 2005Primnoidae Calyptrophora pillsburyae Lesser Antilles 686-1125 Bayer 2001; Cairns 2005Primnoidae Calyptrophora trilepis Straits of Florida, Bahamas 593-911 Bayer 2001; Cairns 2005Primnoidae Candidella imbricata Straits of Florida, Bahamas, Antilles 514-2063

Cairns and Bayer 2004b; Cairns 2005; Watling and Auster 2005; Cairns personal Comm.
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

352

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Isididae Caribisis simplex Western Atlantic >200 Cairns 2005Chrysogorgiidae Chalcogorgia pellucida Western Atlantic >200 Cairns 2005Isididae Chelidonisis aurantiaca mexicana Western Atlantic >200 Cairns 2005Chrysogorgiidae Chrysogorigia agassizii Western Atlantic >200 Cairns 2005Chrysogorgiidae Chrysogorigia desbonni

Greater and Lesser Antilles, Bahamas, Guadeloupe, Martinique, Colombia 155-595

Veronique 1987; Cairns 2001; Cairns 2005; Reyes et al. 2005

Chrysogorgiidae Chrysogorigia elegans

Southeastern Caribbean, Lesser Antilles, Martinique 128-1716 Cairns 2001; Cairns 2005

Chrysogorgiidae Chrysogorigia fewkesii Antilles, Caribbean 403-1200 Cairns 2001; Cairns 2005Chrysogorgiidae Chrysogorigia multiflora Antilles, Bahamas, Straits of Florida 320-1354 Cairns 2001; Cairns 2005Chrysogorgiidae Chrysogorigia spiculosa Gulf of Mexico, Lesser Antilles 914-2256 Cairns 2001; Cairns 2005Chrysogorgiidae Chrysogorigia squamata Antilles, Yucatan 431-1046 Cairns 2001; Cairns 2005Chrysogorgiidae Chrysogorigia thrysiformis Greater and Lesser Antilles, Bahamas 146-526 Cairns 2001; Cairns 2005Coralliidae Corallium medea Straits of Florida >200

Bayer 1961; Cairns 2005; Watling and Auster 2005

Coralliidae Corallium niobe Straits of Florida 659-677

Bayer 1961; Cairns 2005; Watling and Auster 2005

Ellisellidae Ctenocella (Ellisella) atlantica Western Atlantic >200 Cairns 2005Primnoidae Dasystenella acanthina Western Atlantic >200 Cairns 2005Dendrobrachiidae Dendrobrachia multispina Western Atlantic >200 Cairns 2005Anthothelidae Diodogorgia nodulifera?

Throughout Caribbean, including Puerto Rico, Bahamas, Guadeloupe 14-183 Veronique 1987; Bayer 1961; Humann 1993

Chrysogorgiidae Distichogorgia sconsa Straits of Florida >200

Cairns 2005; Watling and Auster 2005

Plexauridae Echinomuricea atlantica Western Atlantic >200 Cairns 2005Ellisellidae Ellisella barbadensis

Striats of Florida, Antilles, including U.S. Virgin Islands 20-488

Bayer 1961; Veronique 1987; Humann 1993; Cairns 2005; Armstrong et al. 2006
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

353

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Ellisellidae Ellisella elongata

Antilles, Puerto Rico, Bahamas, southeastern Caribbean 15-219

Humann 1993; Bayer 1961; Caribbean Fishery Management Council 2004; Cairns 2005

Ellisellidae Ellisella funiculina Western Atlantic >200 Cairns 2005Ellisellidae Ellisella grandiflora Guadeloupe 350 Veronique 1987; Cairns 2005Ellisellidae Ellisella grandis Western Atlantic >200 Cairns 2005Ellisellidae Ellisella nivea Western Atlantic 92~>200 Cairns 2005Ellisellidae Ellisella rosea Western Atlantic 52~>200 Cairns 2005Ellisellidae Ellisella schmitti Western Atlantic 61~>200 Cairns 2005Gorgoniidae Eunicella albatrossae Western Atlantic >200 Cairns 2005Gorgoniidae Eunicella modesta Straits of Florida, Bahamas 274-3236

Watling and Auster 2005; Reed 2006

Gorgoniidae Eunicella tenuis Western Atlantic >200 Cairns 2005Plexauridae Hypnogorgia pendula Guadeloupe, Martinique 176

Veronique 1987; Cairns 2005; Material examined HBOI 26-V-06-1-17 (Reed 2006)

Anthothelidae Iciligorgia schrammi

Straits of Florida, Antilles, southeastern Caribbean 11-366

Humann 1993; Cairns 1977; Cairns 2005

Chrysogorgiidae Iridogorgia pourtalesi Guadeloupe 1343 Cairns 2005; Veronique 1987Isididae Isidella longiflora? Straits of Florida to 667 Reed 2006Ellisellidae Junceella antillarum Western Atlantic >200 Cairns 2005Isididae Keratoisis flexibilis

Straits of Florida, Bahamas, Mexico, Guadeloupe, Colombia, Venezuela 170-878

Cairns 2005; Watling and Auster 2005

Isididae Keratoisis ornata Straits of Florida, Bahamas, Cuba 274-3236

Bedford Institute of Oceanography 2003; Cairns 2005; Watling and Auster 2005

Isididae Keratoisis siemensii Western Atlantic >200 Cairns 2005Isididae Keratosis simplex Martinique 611 Cairns 2005; Veronique 1987Isididae Lepidisis caryophyllia Martinique, Guadeloupe 607-1067 Cairns 2005; Veronique 1987Isididae Lepidisis longiflora Straits of Florida, Caribbean 743-1125

Cairns 2005; Watling and Auster 2005
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

354

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Gorgoniidae Leptogorgia euryale? Southern Lesser Antilles 1060-3000 Bayer 1952; Keller et al. 1975Gorgoniidae Leptogorgia stheno? Southern Lesser Antilles 1060-3000 Bayer 1952; Keller et al. 1975Gorgoniidae Lophogorgia cardinalis Straits of Florida (Cuba) 26-123

Bayer 1961; South Atlantic Fishery Management Council 1998; Cairns 2005

Chrysogorgiidae Metallogorgia splendens Western Atlantic >200 Cairns 2005Plexauridae Muricea laxa Straits of Florida, Bahamas, Antilles 18-128

Bayer 1961; Cairns 1977; Humann 1993; Cairns 2005

Plexauridae Muriceides (=Trachymuricea) hirtaWestern Atlantic >200 Cairns 2005Plexauridae Muriceides kukenthali Western Atlantic >200 Cairns 2005Plexauridae Muriceopsis petila Florida, Bahamas, Guadeloupe 67-455

Cairns 2005; Bayer 1961; Veronique 1987

Primnoidae Narella bellissima

Straits of Florida, Bahamas, Lesser Antilles, including U.S. territories 161-792

Cairns and Bayer 2003; Cairns 2005

Primnoidae Narella pauciflora

Straits of Florida, Antilles, Puerto Rico, Bahamas, Mexico 738-1473

Cairns and Bayer 2003; Cairns 2005

Primnoidae Narella regularis

Straits of Florida, Lesser Antilles and Bahama. 366-792

Veronique 1987; Cairns and Bayer 2003; Cairns 2005; Watling and Auster 2005

Primnoidae Narella spectabilis Only Bahamas. 1485

Cairns and Bayer 2003; Cairns 2005

Primnoidae Narella versluysi Straits of Florida, Cuba. 677-900

Cairns and Bayer 2003; Cairns 2005

Ellisellidae Nicella goreaui Bahamas and southern Caribbean 45-146 Cairns 2007Ellisellidae Nicella americana

Bahamas, Antilles, southern Caribbean 62-237 Cairns 2007

Ellisellidae Nicella deichmannae

Bahamas, northweatern Caribbean, Antilles, incluuding U.S. territories 27-403 Cairns 2007

Ellisellidae Nicella hebes

Western Atlantic, including Bahamas and southern coast of Caribbean to Nicaragua 27-327 Cairns 2005; Cairns 2007

Ellisellidae Nicella lanceolata Only Grand Cayman Island 229-244 Cairns 2007
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

355

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Ellisellidae Nicella obesa

Bahamas, Antilles from Cuba to Barbados, incluuding U.S. territories, Venezuela 174-819

Cairns 2007; Cairns 2005; Veronique 1987

Ellisellidae Nicella guadelupensis

Bahamas, Antilles, including U.S. territories, Venezuela 27-395 Cairns 2007

Ellisellidae Nicella sp. A. Western Atlantic 55-329 Cairns 2007Ellisellidae Nicella gracilis Bahamas, Antilles, Venezuela 60-481 Cairns 2007Ellisellidae Nicella robusta Bahamas, Caribbean 110-259 Cairns 2007Primnoidae Paracalyptrophora carinata Only St. Lucia 514

Cairns and Bayer 2004a; Cairns 2005

Primnoidae Paracalyptrophora duplex

Straits of Florida, Cuba, Antilles, and Bahamas 374-555

Cairns and Bayer 2004a; Cairns 2005

Primnoidae Paracalyptrophora simplex Straits of Florida, Bahamas, Yucatan 165-706

Cairns and Bayer 2004a; Cairns 2005

Paragorgiidae 

Paragorgia (=boschmai)johnsoni Straits of Florida, Bahamas 522-608

Watling and Auster 2005; material examined USNM 35928; Cairns personal comm.

Plexauridae Paramuricea echinata Western Atlantic >200 Cairns 2005Plexauridae Paramuricea grandis Western Atlantic >200 Cairns 2005Plexauridae Paramuricea multispina Western Atlantic >200 Cairns 2005Plexauridae Paramuricea placomus Straits of Florida, Cuba (off Havana) 247-805 Cairns 2005Plexauridae Placogorgia mirabilis Western Atlantic >200 Cairns 2005Plexauridae Placogorgia rudis Western Atlantic >200 Cairns 2005Plexauridae Placogorgia tenius Western Atlantic >200 Cairns 2005; Reyes et al. 2005Plexauridae Placogorgia tribuloidea Western Atlantic >200 Cairns 2005Primnoidae Plumarella aculeata

Bahanas and northern Straits of Florida 400-900

Cairns and Bayer 2004b; Cairns 2005

Primnoidae Plumarella aurea Straits of Florida, Cuba 310-878

Cairns and Bayer 2004b; Cairns 2005

Primnoidae Plumarella dichotoma Straits of Florida 494-1065

Cairns and Bayer 2004b; Cairns 2005
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

356

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Primnoidae Plumarella pellucida Straits of Florida, Bahamas 549-1160

Cairns and Bayer 2004b; Cairns 2005

Primnoidae Plumarella pourtalesii typical Straits of Florida, Cuba 196-882

Cairns and Bayer 2004b; Cairns 2005

Primnoidae

Plumarella pourtalesii var. obtusa Straits of Florida 183-743

Cairns and Bayer 2004b; Cairns 2005; Cairns personal comm.

Primnoidae

Plumarella pourtalesii var. robusta Straits of Florida 183-850

Cairns and Bayer 2004b; Cairns 2005

Primnoidae Primnoa resedaeformis Western Atlantic >200 Cairns 2005Primnoidae Primnoella delicatissima Western Atlantic >200 Cairns 2005Primnoidae Primnoella divaricata Western Atlantic >200 Cairns 2005Primnoidae Primnoella polita Western Atlantic >200 Cairns 2005Isididae Primnosis rigida Western Atlantic >200 Cairns 2005Plexauridae Pseudoplexaura porosa?

Straits of Florida, Antilles, southeastern Caribbean 3-283

Cairns 1977; Kapela and Lasker 1999

Chrysogorgiidae Radicipes gracilis

Straits of Florida, southern Lesser Antilles >200

Keller, et al. 1975; Bedford Institute of Oceanography 2003; Watling and Auster 2005; Cairns 2005

Ellisellidae Riisea paniculata

Bahamas, Caribbean, including U.S. territories 110-704 Cairns 2007

Plexauridae Scleracis guadelupensis Guadeloupe, Martinique 176-350 Veronique 1987; Cairns 2005Plexauridae Scleracis petrosa Guadeloupe, Martinique 275-1607 Veronique 1987; Cairns 2005Isididae Stenisis humilis Western Atlantic >200 Cairns 2005Plexauridae Swiftia casta

Straits of Florida, Cuba (off Havana), Yucatan Channel 40-1953 Cairns 2005

Plexauridae Swiftia exserta

Straits of Florida, Bahamas, Puerto Rico, Mexico, Panama, southern Caribbean 18-494 Veronique 1987; Cairns 2005

Plexauridae Swiftia koreni Straits of Florida, Guadeloupe 221-858 Veronique 1987; Cairns 2005
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

357

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Plexauridae Swiftia pallida (=Thesa nivea) Straits of Florida, Guadeloupe 23-366

Humann 1993; Veronique 1987; Cairns 2005

Plexauridae Swiftia pourtalesii Straits of Florida >200

Cairns 2005; Watling and Auster 2005

Plexauridae Swiftia sp. Sensu Western Atlantic >200 Cairns 2005Keroeididae Thelogorgia longiflora Western Atlantic >200 Cairns 2005Keroeididae

Thelogorgia studeri (=Lingnella richardii) Saint Vincent, Guadeloupe, Barbados 180->200

Veronique 1987; Bayer 1961; Cairns 2005

Keroeididae Thelogorgia vossi Western Atlantic >200 Cairns 2005; Reyes et al. 2005Plexauridae Thesea antiope Western Atlantic >200 Cairns 2005Plexauridae Thesea bicolor Western Atlantic >200 Cairns 2005Plexauridae Thesea gracilis Western Atlantic >200 Cairns 2005Plexauridae Thesea grandiflora Western Atlantic >200 Cairns 2005Plexauridae Thesea grandiflora var. rugulosa Western Atlantic >200 Cairns 2005Plexauridae Thesea grandulosa Western Atlantic >200 Cairns 2005Plexauridae Thesea guadelupensis Guadeloupe >200 Cairns 2005; Veronique 1987Plexauridae Thesea hebes Guadeloupe 275 Cairns 2005; Veronique 1987Plexauridae Thesea parviflora Western Atlantic >200 Cairns 2005Plexauridae Thesea rugosa Western Atlantic >200 Cairns 2005Plexauridae Thesea solitaria Western Atlantic >200 Cairns 2005Plexauridae Thesea sp. Sensu Western Atlantic >200 Cairns 2005Primnoidae Thouarella bipinnata Western Atlantic >200

Cairns 2005; Cairns and Bayer 2006

Primnoidae Thouarella diadema Western Atlantic >200

Cairns 2005; Cairns and Bayer 2006

Primnoidae Thouarella grasshoffi Western Atlantic >200

Cairns 2005; Cairns and Bayer 2006
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

358

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Anthothelidae

Titanideum frauenfeldii (=T. suberosum) Straits of Florida, Cuba 15-238

South Atlantic Fishery Management Council, 1998; Cairns 2005

Chrysogorgiidae Trichogorgia n. sp. Western Atlantic >200 Cairns 2005Plexauridae Villogorgia antillarum Western Atlantic >200 Cairns 2005Plexauridae Villogorgia n. sp. Western Atlantic >200 Cairns 2005Plexauridae Villogorgia nigrescens Martinique, Guadeloupe, Colombia 176-275

Veronique 1987; Cairns 2005; Reyes et al. 2005

Order Helioporacea

Lithotelestidae Epiphaxum micropora Barbados 50-400 Bayer & Muzik 1977, 1979

Order Pennatulacea

Anthoptilidae Anthoptilum grandiflorum Southern Lesser Antilles 1600 Keller et al. 1975Funiculinidae Funiculina quadrangularis Venezuela ~30-4000

Keller et al. 1975; Picton and Howsen 2002

Kophobelemnidae Kophobelemnon irregulatus Southeastern Caribbean Sea Basin 4160 Keller et al. 1975Renillidae Renilla reniformis

Throughout Caribbean, including Puerto Rico to 108 or more

Gosner 1978; Bayer 1961; NOAA 2005

Umbellulidae Umbellula thomsoni

Puerto Rico Trench Caribbean Sea Basin, Cayman Islands 1336-6200 Pasternak 1975

Umbellulidae Umbellula durissima Southeastern Caribbean (Venezuela) ~1012-4400 Pasternak 1975; Keller et al. 1975Umbellulidae Umbellula hemigymna

Southern Lesser Antilles, Caribbean Sea Basin ~2610-3500 Pasternak 1975

Umbellulidae Umbellula magniflora Caribbean Sea Basin ~100-6300 Pasternak 1975; Keller et al. 1975

Class HydrozoaOrder Stylasterina

Stylasteridae Crypthelia glossopoma

Straits of Florida, Yucatan Channel, Lesser Antilles, including U.S. territories 198-864 Cairns 1986

Stylasteridae Crypthelia insolita Southernmost Lesser Antilles 159-720 Cairns 1986Stylasteridae Crypthelia papillosa Lesser Antilles 161-545 Cairns 1986
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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359

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Stylasteridae Crypthelia peircei

Greater and Lesser Antilles, including U.S. territories 159-837 Cairns 1986

Stylasteridae Crypthelia tenuisepata

Lesser Antilles, including U.S. territories 761-1061 Cairns 1986

Stylasteridae Distichopora anomala Lesser Antilles 139-311 Cairns 1986Stylasteridae Distichopora barbadensis Southernmost Lesser Antilles 102-311 Cairns 1986Stylasteridae Distichopora cervina Lesser Antilles, Puerto Rico 68-384 Cairns 1986Stylasteridae Distichopora contorta Only off Havana 125-368 Cairns 1986Stylasteridae Distichopora foliacea Straits of Florida, Yucatan Channel 183-366 Cairns 1986Stylasteridae Distichopora rosalindae Western Caribbean 165-198 Cairns 1986Stylasteridae Distichopora sulcata Only off Havana, Cuba 1097 Cairns 1986Stylasteridae Distichopora uniserialis Only off Havana, Cuba 333-366 Cairns 1986Stylasteridae Distichopora yucatanensis Western Caribbean 39-261 Cairns 1986Stylasteridae Errina altispina Only Yucatan Channel 198-309 Cairns 1986Stylasteridae Errina cochleata Straits of Florida, Bahamas 194-534 Cairns 1986Stylasteridae Lepidopora biserialis Straits of Florida 196-370 Cairns 1986Stylasteridae Lepidopora carinata Only off Havana 60-494 Cairns 1986Stylasteridae Lepidopora clavigera Only off Barbados 150-282 Cairns 1986Stylasteridae Lepidopora decipiens Lesser Antilles 270-670 Cairns 1986Stylasteridae Lepidopora glabra Only off Havana 267-1170 Cairns 1986Stylasteridae Lepidotheca brochi

Dominica, Martinique, Montserrat, Saint Kitts and Nevis 545-864 Cairns 1986

Stylasteridae Lepidotheca pourtalesi Straits of Florida 123-368 Cairns 1986Stylasteridae Pliobothrus echinatus

Lesser Antilles and Mona Passage, including Puerto Rico 164-708 Cairns 1986

Stylasteridae Pliobothrus symmetricus

Throughout Caribbean, including Puerto Rico 150-400 Cairns 1986

Stylasteridae Pliobothrus tubulatus Antilles, including U.S. territories 419-708 Cairns 1986Stylasteridae Stenohelia pauciseptata Only off St. Lucia 300-514 Cairns 1986
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

360

Higher 

Taxon Family Species ▲ Distribution ♠ Depth Range Reference ‡

Stylasteridae Stylaster antillarum

Lesser Antilles and the Mona Passage, including Puerto Rico 174-653 Cairns 1986

Stylasteridae Stylaster atlanticus

Only off Puerto Rico (Isla de Culebra) 823 Cairns 1986

Stylasteridae Stylaster auranticus Greater Antilles, off Cuba 112-377 Cairns 1986Stylasteridae Stylaster complanatus

Straits of Florida, Antilles, includingU.S. territories 183-707 Cairns 1986

Stylasteridae Stylaster corallium

Barbados, Dominica, Grenada, Martinique, Saint Lucia 13-298 Cairns 1986

Stylasteridae Stylaster duchassaingi

Widespread Caribbean, including U.S. Virgin Islands, Bahamas

42-692 (common 

200-400) Cairns 1986

Stylasteridae Stylaster erubescens

Continental slopes of the southeasetern United States to the Yucatan Peninsula 146-965 Cairns 1986

Stylasteridae Stylaster filogranus Straits of Florida 183-274 Cairns 1986Stylasteridae Stylaster inornatus Only off the Yucatan Peninsula 198-309 Cairns 1986Stylasteridae Stylaster laevigatus Straits of Florida, Yucatan Channel

123-759 (common 

300-400) Cairns 1986

Stylasteridae Stylaster miniatus Straits of Florida 146-530 Cairns 1986Stylasteridae Stylaster profunda Lesser Antilles, including Puerto Rico

159-2021 

(common 200-650)Cairns 1986

Stylasteridae Stylaster roseus

Widespread Caribbean, including Puerto Rico

0.5-373 (common 

0.5-30) Cairns 1986

Stylasteridae Stylaster spatula Only off southeastern Puerto Rico 384-549 Cairns 1986
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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361

Appendix 8.2. An aggregate inventory of the benthos associated with the major structure-forming corals E. profunda, L. pertusa, M. carolina, M. oculata, and S. varibilis throughout the wider Caribbean. Benthos is grouped by coral species. Sources include station data from 28 locations in the wider Caribbean, sampled by the following research vessels: R/V Pillsbury (14 Stations); R/V Columbus Iselin (5 Stations); R/V Gerda (6 Stations); R/V Gilliss (1 Station); Submersible Alvin (multiple station); and R/V Eastward (multiple stations) (Anon 1963; 1964, 1965, 1966, 1968, 1969, 1970, 1972a, 1972b, 1973, 1974; Bayer 1966; Staiger 1968a, 1968b, 1969, 1971; Voss 1966a and 1966b). Locations are identified in Figure 3 by the letter S (some station locations overlap). *Please note that many species noted were first identifications and spelling is presented as it appears on the original station sorting sheets.

Potential 

structure and 

habitat- forming 

species

Other 

coelenterata

Crustacean (deep-seas 

crabs, shrimp and 

lobsters)

Fish Echinoderms Mollusks Other

E. profundaL. pertusa;S. varibilis; gorgonians; hydroids; octocorals

"Solitary corals"; "Medusae"

Homolodramia paradoxa; Galatheid sp.; Pylocheles scutata; Glyphocrangon longleyi; Nephropsis rosea; Plesionika acanthonotus; Pasiphaeidae sp.; Stylirostrissp.; Nematocarcinua cursor; isopods

Benthobatis sp.; Breviraja plutonia (ray); Cruriraja sp. (skates); Etmopterus sp. (dogfish sharks); Laemonema sp. (hake); Synaphobranchus sp.; Nettastomatidae sp. (eels)

Gastropods; pelecypods; scaphopods; brachiopods

Polychaetes; porifera; pyncnogonids

L. pertusaS. varibilis; E. profunda; gorgonians; "corals"

"Solitary corals"; actinarians

Aristaeomorphasp.; Bathyplax sp.; Benthochascon sp.; Crangonidae sp.; Galatheidsp.; Hymenopenaeus sp.; Munida sp.; Munidopsis sp.; Nephropsis sp.; Palicus sp.; Phoberus sp.; Polychelessp.; Raninoides sp.; Rochiniasp.; Stereomastis sp.; Stylirostris sp.; Xanthidae sp.

Bathyclupea sp. (deepsea herring); Bembrops sp.; Benthobatis sp.; Breviraja plutonia (ray); Brotulidsp. (fangtooths); Chaunax pictus(anglerfish); Chlorophthalmus sp.; Cruriraja sp. (skates); Dibranchussp. (batfishes); Etmopterus sp. (dogfish shark); Gadidae sp (cod) ; Galeus sp. (benthic sharks); Laemonema sp. (hake); Macrouridsp. (grenadiers); Merluccius sp. (hake); Moridae sp. (deepsea codfishes); Neoscopelus sp. (lanternfish); Nettastomidsp.; Promyllanter schmitti; Synaphobranchus sp.

Holothurians (sea cucumbers); Nymphaster sp.; Ophiuroids; Phormosoma sp.; Zoroaster sp.

Arca sp.; Columbarium sp.; Gastropods; Gaza sp.; Natica sp.; Turridae sp.; Voluta sp.; Xenophora sp.

Polychaetes; porifera
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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362

Potential 

structure and 

habitat- forming 

species

Other 

coelenterata

Crustacean (deep-seas 

crabs, shrimp and 

lobsters)

Fish Echinoderms Mollusks Other

M. carolina"Corals"; Antipatharia sp. (black corals); Stylasterina sp. (hydrocorals)

"Solitary corals"; Actiniaria sp. (anemones); Alcyonaria sp. (soft corals); Hydroids; Urochordatasp. (encrusting gorgonians); zoanthids

Anomalothir furcillatus; Brachyura sp.; Cancellus oratus; Cirripedia sp.; Clibanarius anomalus;; Galatheid sp.; Homaridaesp.; Iliacantha subglobosa; Leucosiidae sp.; Majidaesp.; Munida irrasa; Munida schroederi ; Myropsis quinquespinosa; Nephropsis aculeate; Paguristes spinipes; Pandalidae sp.; Plesionika acanthonotus; Podochela curvirostris ; Pyromia cuspidate; Sicyonia stimpsoni; stomatopods; Sympagurus pictus

Calamopteryx goslinei; elasmobranch egg cases

Asteroidea; crinoids; echinoids; holothurians (sea cucumbers); ophiuroids; starfish; brittle stars; urchins

Gastropods; pelecypods; brachiopods

Algae; "Large sponge 18" across"; oolite chunk; polychaetes; bryozoans; tunicates
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

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363

Potential 

structure and 

habitat- forming 

species

Other 

coelenterata

Crustacean (deep-seas 

crabs, shrimp and 

lobsters)

Fish Echinoderms Mollusks Other

M. oculataS. varibilis; "pink gorgonians"; "various coelenterates"; Acanella sp. (bamboo corals); Bathypathes patula; Calytrophora pourtalesi (sea fan / sea whip); Chrysogorgia sp. (sea fan / sea whips); Cirripathessp. (sea whips); gorgonians; Iridogorgia sp. (spiral corals); Keratoisis sp.; Plumarella sp. 

"Solitary corals"; Deltocyathus sp.; Desmophyllum sp.; Hydroids; Odontocyathus coronatus; Pennatulidae sp. (sea pens); Stephanocyathus diadema; Telestula sp. (anemones); aoanthids

"Various crustaceans"; Acanthephyra sp.; Amphipods; Aristaeinaesp.; Aristeus sp.; Axiidaesp.; Bathymunida sp.; Bathypalaemonella sp.; Bathyplax sp.; Bathysquillasp.; Galatheid sp.; Gennadassp.; Glyphocrangon sp.; Gnathophausia sp.; Heterocarpus sp.; isopods; Lithodid sp.; Majidae sp.; Munida sp.; Munidopsissp.; Nematocarcinussp.; Nephropsis sp.; Oplophorus sp.; Pandalidaesp.; Parapagurus sp.; Plesiopenaeus sp.; Polycheles crucifer; Polychelidae sp.; Pontophilussp.; Prionocrangon sp.; Scalpellum sp.; Sergestessp.; Spongicoloidessp.; Stereomastis sp.; Stylodactylus sp.; Uroptychusnitidus; Uroptychussp; Uroptychus sp.; Verrucomorphs sp.

Acanthurus sp. (surgeonfish); Alepocephalus sp.; Apristurussp. (cat sharks); Barathronussp. (cusk-eels); Bathypterois sp. (tripod fish); Benthobatis sp.; Brotulid sp. (fangtooths); Chaunax pictus (anglerfish); Diaphussp. (headlightfish); Dibranchus atlanticus (Atlantic batfish) ; Dibranchus sp. (batfishes); Dicrolene intronigra; Diretmus sp. (dorys); Gonostomatidae sp. (bristlemouth); Halosaurs sp.; Hariotta sp. (chimaera); Ilyophis sp. (eels); Macrourid sp. (grenadiers); Monomitopus sp. (cusk-eels); Myctophid sp. (lanternfish ); Myxini sp. (hagfish); Neoscopelus sp. (lanternfish); Nesiarchus sp. (scabbardfish); Notacanthid sp. (spiny eel); Oxygadus sp. (rattails); Promyllantor schmitti; Saurida sp. (lizardfish); Searsid sp.; Sternoptychidae sp. (hatchetfish); Stomias sp. (dragon fish ); Synaphobranchus oregoni Synaphobranchus sp.

"Pancake urchin"; "Soft urchin"; asteroids; Benthopectinids; Crinoids; Goniaster sp.; holothurians (sea cucumbers); Molpadia sp. (sea cucumbers); Nymphaster sp.; ophuiroids; Phormosoma sp. (sea biscuts); Psilaster squameus; Zoroaster fulgens

"Squid, unidentified"; Acesta sp. (Clams); Arenatus sp.; Gaza superba; Leptothyra sp.; Leucosyrinxsp.; Limopsissp.; Melongenasp; Mitra sp.; Pseudamusium dallil (bivalve, clam?); scaphopods; Solariella sp.; solenogaster (shelless mollusk); Spirula spirula; Turridaesp.

"Grapefruit sponges"; Cladorhizasp (potato sponges); Hexactinellidasp. (glass sponges); Hyalonemasp.; polychaetes
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

364

Potential 

structure and 

habitat- forming 

species

Other 

coelenterata

Crustacean (deep-seas 

crabs, shrimp and 

lobsters)

Fish Echinoderms Mollusks Other

S. varibilisM. oculata; L. pertusa; E. profunda; "corals";Acanella sp. (bamboo corals);Callogorgia sp.;Chrysogorgia sp. (sea fan / sea whips); Keratoisis sp. (bamboo corals); Nicella sp.; octocorals;Primnosis sp. 

"Solitary corals"; Odontocyathus coronatus; Stephanocyathus diadema; Telestula sp. (anemones); Clavulariidae sp.

Acanthephyra sp.; amphipods; Aristaeinae sp.; Axiidae sp.; Chirostylus sp.; Galatheid sp.; Gennadassp.; Glyphocrangon aculeate; Glyphocrangon alispina; Glyphocrangon sp.; Heterocarpus sp.; isopods; Lithodid sp.; Majidae sp.; Mixtopagurus paradoxus; Munida sp.; Munidopsissp.; Nematocarcinus sp.; Nephropsis sp.; Oplophorussp.; Paguristes sp.; Penaeid sp.; Polycheles crucifer; Polychelidaesp.; Prionocrangon sp.; Processa sp.; Pylopagurussp; Rochinia sp.; Sergestessp.; Spongicoloidessp.; Stereomastis sp.; Stylodactylus sp.; Systellaspis sp.; Uroptychussp.

Brotulid sp. (fangtooths); Callionymus sp.; Chaunax sp. (Anglerfish); Chlorophthalmussp.; Dibranchus atlanticus(Atlantic batfish); Gadomus longifilis (Bathygadid family); Hymenocephalus sp. (rattails); Macrourid sp. (grenadiers); Monomitopus sp. (cusk-eels); Peristedion sp. (sea robins); Promyllantor schmitti; Saurida sp. (lizardfish)

Ceramaster elongatus(Cookie starfish); Ceramastersp.; cidarids; Circeaster americanus; Lophaster verrilli; Nymphastersp.; Ophidiastersp.; Solaster caribbea; Spatangoids; Zoroaster ackleyi

"Finned octopod"; "unidentified squid"; Acestasp. (Clams); brachiopods; Dentalium sp.; Naticidae sp.; Ornithoteuthis antillarum ; Propeamusium dalli; pteropods; scaphopods; Solariella sp.; Solenogastersp.; Turridae sp.

Cladorhiza sp. (potato sponges); polychaetes; pyncnogonids
STATE OF DEEP CORAL ECOSYSTEMS IN THE CARIBBEAN REGION

CARIBBEAN

365

Species Depth (m)

Snappers:

Apsilus dentatus (black snapper) 70 – 290

Etelis oculatus (queen or green snapper) 180 – 463

Lutjanus buccanella (blackfin snapper) 80 – 180

L. vivanus (silk snapper) 60 – 210

Rhomboplites aurorubens (vermilion snapper) 80 – 230

Sharks:

Centrophorus granulosus (gulper shark) to 210

Eugomphodus taurus (grey nurse shark) to 210

Hexanchus vitulus (bigeyed sixgill shark) to 250

Mustelus canis (smooth dogfish) to 250

Scyliorhinus boa (catshark) to 270

Sphyma lewini (scalloped hammerhead shark) to 170

Squalus cubensis (Cuban dogfish) to 300

Promthicthys Prometheus (rabbit-fish) to 170

Groupers & hinds:

Epinephelus adscensionis (rock hind) 0 – 120

E. flavolimbatus (yellowedge grouper) to 132

E. fulvus (coney) 0 – 230

E. guttatus (red hind) 0 – 200

E. morio (red grouper) to 174

E. mystacinus (misty grouper) 180 – 270

Mycteroperca phenax (scamp) to 174

Neoscombrops sp. to 190

Other:

Erythrocles monody (Atlantic rubyfish) to 300

Gephyroberix darwini to 310

Ostichthys trachypoma (bigeye soldierfish) to 290

Polymixia lowel (beardfish) to 260

Pristipomoides sp. (jobfish) 180 – 300+

Appendix 8.3. Commercially important deep-sea fish of the wider Caribbean 

(FAO 1993).

United States Department of Commerce

Carlos M. Gutierrez

Secretary

National Oceanic and Atmospheric Administration

Vice Admiral Conrad C. Lautenbacher, Jr. USN (Ret.)

Under Secretary of Commerce for Oceans and Atmosphere

National Marine Fisheries Service

Dr. William T. Hogarth

Assistant Administrator for Fisheries