‐ ‐

The State of Deep‐Sea Coral and Sponge

Ecosystems of the United States

Thomas F. Hourigan, Peter J. Etnoyer, and Stephen D. Cairns

U.S. Department of Commerce

National Oceanic and Atmospheric Administration

National Marine Fisheries Service

NOAA Technical Memorandum NMFS OHC 4

December 2017

                 

     

                   

                     

                 

                     

                       

                   

     

   

     

           

     

               

  

       

           

The State of Deep‐Sea Coral and Sponge Ecosystems of

the United States

Thomas F. Hourigan1, Peter J. Etnoyer2, and Stephen D. Cairns3

1 NOAA, National Marine Fisheries Service, Office of Habitat Conservation, Deep Sea

Coral Research and Technology Program, Silver Spring, MD 20910

2 NOAA, National Ocean Service, National Centers for Coastal Ocean Science, Center

for Coastal Monitoring and Assessment, 219 Fort Johnson Rd., Charleston, SC 29412

3 Smithsonian Institution, National Museum of Natural History, P.O. Box 37012,

Washington D.C., 20013

NOAA Technical Memorandum NMFS‐OHC‐4

December 2017

U.S. Department of Commerce

Wilbur L. Ross, Jr., Secretary

National Oceanic and Atmospheric Administration

RDML Tim Gallaudet, Ph.D., USN Ret., Acting NOAA Administrator

National Marine Fisheries Service

Chris Oliver, Assistant Administrator for Fisheries
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Recommended citation for full report:

Hourigan TF, Etnoyer PJ, Cairns SD (2017) The State of Deep‐Sea Coral and Sponge

Ecosystems of the United States. NOAA Technical Memorandum NMFS‐OHC‐4. Silver

Spring, MD. 467 p.

Recommended citation for individual chapters (e.g., Alaska Chapter):

Stone RP, Rooper CN (2017) State of Deep‐Sea Coral and Sponge Ecosystems in the

Alaska Region. In: Hourigan TF, Etnoyer PJ, Cairns SD (eds.). The State of Deep‐Sea

Coral and Sponge Ecosystems of the United States. NOAA Technical Memorandum

NMFS‐OHC‐4. Silver Spring, MD. 35 p.

Copies of this report may be obtained from:

Heather Coleman

NMFS Office of Habitat Conservation

National Oceanic and Atmospheric Administration

1315 East‐West Highway, Room 14201

Silver Spring, MD 20910

Or online at:

http://deepseacoraldata.noaa.gov/library or http://spo.nmfs.noaa.gov/tm/

Cover photos: Top – A sculpin rests on a large red tree coral off Alaskaʹs Glacier Bay National

Park. Image courtesy of the Deepwater Exploration of Glacier Bay National Park expedition and

University of Connecticut Northeast Underwater Research, Technology & Education Center.

Bottom – View of remotely operated vehicle Deep Discoverer investigating a large primnoid coral

colony. Image courtesy of the NOAA Office of Ocean Exploration and Research.
iii

   

Table of Contents 

Executive Summary..................................................................................................................... v

Acknowledgments ...................................................................................................................... xi

1. Introduction to the State of Deep‐Sea Coral and Sponge Ecosystems of the U.S........... 1

2. Species Discovery of Deep‐Water Corals and Sponges in U.S. Waters (2007‐2016) .... 39

3.              State of Deep‐Sea Coral and Sponge Ecosystems          of the Alaska Region........................    57

4. Effects of Fishing Gear on Deep‐Sea Corals and Sponges in U.S. Waters ..................... 93

5. State of Deep‐Sea Coral and Sponge Ecosystems of the U.S. West Coast ................... 113

6. An Update on Recent Research and Management of Hawaiian Black Corals............ 157

7. State of Deep‐Sea Coral and Sponge Ecosystems of the U.S. Pacific Islands Region 173

8. Predictive Habitat Modeling for Deep‐Sea Corals in U.S. Waters................................ 213

9. State of Deep‐Sea Coral and Sponge Ecosystems of the Northeast United States ..... 237

10. Age, Growth Rates, and Paleoclimate Studies in Deep‐Sea Corals of the U.S.......... 299

11. State of Deep‐Sea Coral and Sponge Ecosystems of the Gulf of Mexico Region...... 321

12. Population Connectivity of Deep‐Sea Corals................................................................. 379

13. State of Deep‐Sea Coral and Sponge Ecosystems of the Southeast United States.... 409
     

       

           

     

Zoanthids overgrowing coral off

the Northwest Hawaiian Islands.

Courtesy of the NOAA Office of

Ocean Exploration and Research.
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Executive Summary 

Corals and sponges create the most

important biogenic habitats in the deep sea,

and support ecosystems of incredible

variety and biodiversity. In 2007, the United

States National Oceanic and Atmospheric

Administration (NOAA) published the first

peer‐reviewed report on the State of Deep

Coral Ecosystems of the United States

(Lumsden et al. 2007). The 2017 report on

the State of Deep‐Sea Coral and Sponge

Ecosystems of the United States updates

information on deep‐sea coral ecosystems

and management efforts to protect them

over the last decade, and presents a first

summary of information on U.S. deep‐sea

sponge ecosystems. It consists of an

introduction, six regional chapters with

accompanying on‐line resources, and six

spotlight chapters that highlight advances

on crosscutting themes.

The introductory chapter (Hourigan et al.,

Chapter 1) outlines the purpose of the

report and introduces deep‐sea coral and

sponge ecosystems. It presents national‐

level efforts since 2007 that have influenced

regional‐level research and conservation.

NOAA’s 2010 Strategic Plan for Deep‐Sea

Coral and Sponge Ecosystems: Research,

Management, and International Cooperation

guides the agency’s approach. New

research on these ecosystems has been led

by NOAA’s Deep Sea Coral Research and

Technology Program, which began

operations in 2009, and by NOAA’s Office

of Ocean Exploration and Research in

partnership with the Department of the

Interior’s Bureau of Ocean Energy

Management (BOEM) and U.S. Geological

Survey (USGS). Chapter 1 concludes with a

synthesis of the conservation status of U.S.

deep‐sea coral and sponge ecosystems as

revealed by the regional chapters that

follow.

Cairns et al. (Chapter 2) present a brief

introduction to discovery of deep‐sea coral

and sponge species. As of 2016, 662 species

of corals are known to occur in U.S. waters

below 50 m. This list includes 62 new

species and three new genera described

since 2007. Octocorals (soft corals,

gorgonians and sea pens) account for over

half of deep‐sea coral species richness,

followed by the azooxanthellate Scleractinia

(stony corals), Antipatharia (black corals),

and the stylasterids (lace corals). The Gulf

of Mexico has been the most extensively

sampled region for deepwater corals and

has the highest recorded species richness

(227 species), but it is clear that a large

number of undescribed species still occur in

each U.S. region. In contrast to corals, the

deep‐water sponge fauna of the U.S. has not

been systematically inventoried. It is likely

that the species richness of deepwater

sponges exceeds that of corals, and research

over the last decade has resulted in many

new collections in Alaska, the West Coast

and Hawai‘i. However most specimens

remain to be described. New genetic

techniques are complementing existing

morphological analyses, and hold promise

of expanding our knowledge of both the

taxonomy and population genetics of deep‐
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sea corals and sponges. The first ever

Comprehensive List of the Deep‐Water Corals

occurring in the EEZ of the United States and

its Possessions is an online supplement that

documents all deepwater coral species

reported to occur within U.S. waters as of

2016. The comprehensive list is derived

from seven regional species lists, all of

which are available on NOAA’s Deep‐Sea

Coral Data Portal:

https://deepseacoraldata.noaa.gov/library/2

017‐state‐of‐deep‐sea‐corals‐report.

Stone & Rooper (Chapter 3) report on

research conducted in Alaska since 2007,

including extensive new surveys under

NOAA’s Deep Sea Coral Research &

Technology Program. This research

continues to show that the region is home to

diverse ecosystems supported by

extraordinary coral and sponge resources.

Of particular importance are the rich deep‐

sea coral and sponge gardens in the

Aleutian Islands and the red‐tree coral

groves in the Gulf of Alaska, both of which

are more extensive than previously known.

These aggregations appear to provide

important habitat to several fisheries of

national and international importance.

Bottom‐contact fisheries continue to be the

primary threat to these habitats, and

bycatch of corals and sponges is high

compared to other U.S. regions, especially

in certain Aleutian Islands bottom trawl

fisheries.

Bottom‐contact fishing, particularly bottom

trawling, is the most immediate threat to

deep‐sea coral and sponge habitats in most

areas where such fisheries are active.

Rooper et al. (Chapter 4) summarize the

known interactions between fishing gear

and deep‐sea corals and sponges in U.S.

waters. They highlight studies on the effects

of mobile and fixed‐location fishing gears

on deep‐sea corals and sponges. Damage

from mobile gear (e.g., bottom trawls) is

well documented. Fewer studies have

explored impacts of fixed gears, such as

bottom‐set longlines, gillnets and traps.

These deserve additional attention, as they

may be used in rougher habitats that are

preferred by deep‐sea corals and sponges,

but are inaccessible to trawls. Many

vulnerable deepwater areas in the U.S.

exclusive economic zone (EEZ) have been

closed to gears that impact deep‐sea corals

and sponges – considered the most effective

management measure to address fisheries

interactions. There has also been some

success in modifying fishing gears and

practices (e.g., reducing roller size or raising

the sweeps on trawls) that may reduce

impacts of these gears on deep‐sea corals

and sponges.

Clark et al. (Chapter 5) review information

since 2007 that has advanced our

understanding of deep‐sea coral and sponge

abundance and distribution off the U.S.

west coast. This understanding has been

catalyzed by new research under NOAA’s

Deep Sea Coral Research & Technology

Program and data syntheses associated with

the Pacific Fishery Management Council’s

5‐year review of Pacific coast groundfish
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essential fish habitat (EFH). Deep‐sea corals

on hard substrata are widely distributed

and the composition of assemblages

changes with depth. The dominant species

in many areas are small and occur at

relatively low densities, but in certain areas

(e.g., ridges of seamount and certain banks,

particularly in the Southern California

Bight), large gorgonians may occur in

patches of high density. As in other regions,

information on deep‐sea sponges is much

more rudimentary, however, it appears that

in many hard‐bottom habitats in deeper

waters, sponges are more abundant than

corals.

Fishing with bottom‐contact gear, especially

bottom trawls, remains the most immediate

threat to both deep‐sea coral and sponge

ecosystems. While many areas expected to

be prime habitat for corals and sponges

were protected from trawling either

indirectly in 2001 or directly in 2006,

continued bycatch in commercial trawl

fisheries indicates that significant

interactions remain, and in many cases are

restricted to relatively discrete areas.

Among other stressors, ocean acidification

will be a major long‐term concern for some

corals. Since 2007, new protected areas have

been established in state waters, and

National Marine Sanctuaries (Monterey Bay

and the Greater Farallones) have been

expanded to include deeper habitats

containing important deep‐sea coral and

sponge ecosystems.

Hawai‘i is the only U.S. state with major

black and precious coral harvest industries.

Wagner et al. (Chapter 6) provide a brief

history of the black coral fishery and its

current management. They show how

recent research on the taxonomy,

distribution, reproduction and life history

of black corals has increased our knowledge

of these taxa and may inform future

management.

The U.S. Pacific Island Region consists of

more than 50 oceanic islands stretching over

a vast area of the central and western

Pacific. Parrish et al. (Chapter 7) summarize

new information on the deep‐sea coral and

sponge ecosystems found on island slopes,

oceanic ridges and seamounts of the region.

As in the 2007 report, most of the

information from the region is limited to the

Hawaiian Archipelago, however, a major

three‐year NOAA campaign of exploration

and research is underway, which will

provide the first surveys of many deep‐sea

areas in the U.S. territories. The last ten

years of research have used available data

to study the taxonomy, biology,

reproduction and growth of certain coral

taxa – especially black corals. Habitat

suitability models have been developed for

deepwater corals, and the first steps to

characterize the sponge community has

begun. Hawai‘i’s deep‐sea sponges

represent a rich and diverse fauna that form

important, and sometimes extensive,

biogenic habitats.
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In 2009, major areas of the U.S. Pacific

Islands were included in three new Marine

National Monuments (Marianas Trench,

Rose Atoll, and Pacific Remote Islands

Marine National Monuments [PRIMNM]).

Portions of the PRIMNM were expanded in

2014, and the Papahānaumokuākea Marine

National Monument was extended to

encompass the entire EEZ around the

Northwestern Hawaiian Islands in 2016,

forming the largest contiguous fully

protected conservation area in the U.S., and

currently the largest marine conservation

area in the world.

Predictive habitat modeling is increasingly

used to help identify areas that are most

likely to harbor deep‐sea coral and sponge

taxa. Guinotte et al. (Chapter 8) review

modeling methodologies and their

application in each U.S. region. The growth

and adoption of these techniques are fueled

by recent improvements in input data

quality/quantity, the low cost of producing

and updating the models, and the need to

identify habitat across large spatial extents

for management. The spatial resolution of

model results has improved in the last

decade to the point where model outputs

are now used to target areas for field

sampling efforts and to help inform

regional Fishery Management Council

management actions designed to protect

deep‐sea coral habitats. Both the North

Pacific and Mid‐Atlantic Fishery

Management Councils have recently used

predictive deep‐sea coral habitat models to

inform management decisions.

Packer et al. (Chapter 9) review research

and management of deep‐sea corals and

sponges in the U.S. Northeast region since

2007. Multibeam mapping and initial visual

surveys since 2010 have been conducted in

almost all the submarine canyons off the

Northeast coast, as well as the New

England seamounts within the EEZ, and

significant areas in the Gulf of Maine. Rich

octocoral gardens were discovered at

several sites within the Gulf of Maine that

serve as habitat for large numbers of

redfish. Studies in Baltimore and Norfolk

Canyons off Virginia allowed more detailed

descriptions of the distribution and life

history of deep‐sea corals. Knowledge gaps

still exist for many of these organisms,

especially for sponges, but these studies

have contributed greatly to our regional

knowledge of deep‐sea coral diversity,

distribution, and habitat characteristics.

Based on this new information, and

recognizing that bottom‐contact fisheries

represent the most urgent threat to these

habitats, the New England and Mid‐

Atlantic Fishery Management Councils

have moved forward on plans for deep‐sea

coral habitat protection, especially those for

the submarine canyons and the Gulf of

Maine. This action led to the creation of the

99,000 km2 Frank R. Lautenberg Deep‐Sea

Coral Protection Area in the Mid‐Atlantic,

and was a major factor behind creation of

the Northeast Canyons and Seamounts

Marine National Monument. The New

England Fishery Management Council has

proposed new deep‐sea coral protections
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for the Gulf of Maine, and is working on

additional protections for offshore canyon

areas in 2017.

Deep‐sea corals are among the slowest

growing, longest‐lived skeletal accreting

marine organisms. Prouty et al. (Chapter

10) explore the new techniques that are

yielding information about the age and

growth of deep‐sea corals. Studies over the

last decade have revealed black and gold

corals that can live for thousands of years.

Understanding age, growth rate, and

lifespan characteristics allows assessment of

vulnerability and recovery from

perturbations for coral habitats, and can

thereby inform effective management and

conservation strategies. The ability to

accurately age deep‐sea corals has allowed

their use as biogeochemical proxies that

provide a unique view of marine climate

and environmental change over time. This

progress has allowed scientists to look back

into the coral’s climate history using stable

and radioisotope techniques, and

incorporate this information into models of

future climate change.

As described by Boland et al. (Chapter 11),

the deep‐sea coral habitats of the Northern

Gulf of Mexico have received the most

extensive research of any U.S. region over

the last decade. This coverage includes

major collaborative research programs led

by BOEM, ongoing research by NOAA’s

Flower Garden Banks National Marine

Sanctuary, as well as natural resource

damage assessments following the

Deepwater Horizon oil spill. The

availability and interpretation of industry‐

provided seismic reflectivity data have

enhanced our understanding of the

relatively rare areas of hard substrata that

define the distribution of most of the

region’s deep‐sea corals. Studies have led to

increased understanding of genetic

diversity, age, growth, and reproduction of

deep‐sea corals. Sponge communities (at

least high‐density communities) appear to

be relatively rare in the Gulf of Mexico

compared to corals.

The Deepwater Horizon incident was a

landmark event with major revelations

related to deep spill impacts including the

dynamics of deep plumes, and potential for

impacts to deepwater habitats from

horizontal transport of hydrocarbon plumes

and from oiled marine snow. Octocoral

habitats at both deep‐sea and mesophotic

depths were among the habitats damaged

in the spill. The results of research and

damage assessments have led to increased

awareness among managers and the public.

BOEM has instituted new regulatory

policies for the oil and gas industry dealing

with biological communities in the Gulf, in

addition to numerous changes in regulatory

policies related to drilling safety. The

Flower Garden Banks National Marine

Sanctuary is currently exploring alternatives

for boundary expansion to protect known

high value benthic habitats and cultural

resources in the north‐central Gulf of

Mexico.
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Morrison et al. (Chapter 12) review research

on population connectivity of deep‐sea

corals. Understanding connectivity through

exchange of larvae among populations and

the processes that influence connectivity can

help inform conservation efforts. The

authors provide an overview of concepts

and discuss how new molecular genetic

techniques are being used to understand the

population structure of deep‐sea corals.

They review a number of studies in U.S.

waters, including the first ocean basin‐scale

description of the population structure of

the most important reef‐forming deep‐sea

coral, Lophelia pertusa. They conclude by

identifying specific research questions that

may guide future studies of connectivity

among deep‐sea corals.

Hourigan et al. (Chapter 13) review how

new research has improved our

understanding of deep‐sea coral and sponge

ecosystems of the Southeast U.S., and how

this new understanding has informed major

new management efforts. New mapping

and surveys have confirmed that the

Southeast U.S. harbors the nation’s highest

concentration of deep‐sea coral reefs, and

probably ranks among the top such

provinces globally. In addition to branching

stony corals, gorgonians, black corals and

sponges are important contributors to

habitat structure in areas of hard bottoms,

enhancing local biodiversity by providing

habitat for a large number of associated

species. Despite research progress, much of

this region remains unexplored. The last

decade has also seen regional‐scale actions

to protect deep‐sea coral reef provinces

from fishing impacts through a series of

deepwater Coral Habitat Areas of Particular

Concern. A new network of shallower

marine protected areas is providing similar

benefits to shelf‐edge coral and sponge

habitats. These new fishing measures,

developed by the South Atlantic Fishery

Management Council and implemented by

NOAA, have significantly reduced the

scope for fishing damage to deep‐sea coral

and sponge habitats.

Thank you for reading. We look forward to

the next ten years of deep‐sea exploration

and discovery.
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Acknowledgments 

This report would not have been possible without the assistance of Heather Coleman, Catherine

Polk, and Fan Tsao, who contributed many hours to layout and organization of the report.

Robert McGuinn was responsible for all of the annex coral maps and many of the individual

chapter maps. The editors are indebted to the large number of regional and subject matter

experts who served as peer reviewers for each chapter and online species list. Special thanks to

Andrea Quattrini, Sandra Brooke, Joshua DeMello, and Les Watling, who reviewed several

chapters; and Marcelo Kitahara, Alberto Lindner, Dennis Opresko, Andrea Quattrini, Daniel

Wagner, and Gary Williams, who reviewed multiple regional coral species lists. Finally, we

wish to thank all of the authors for their patience during the extended development time for

this report.

Unless otherwise noted, photographs are credited to NOAA. Most images were taken during

expeditions sponsored by NOAA’s Office of Ocean Exploration and Research and NOAA’s

Deep Sea Coral Research and Technology Program.
xii

Iridogorgia soft coral with 

squat lobsters in the 

northwestern Gulf of 

Mexico. Courtesy of the 

NOAA Office of Ocean 

Exploration and Research.
                           

     

           

                   

                 

                 

                       

                     

                 

               

                 

                   

                     

               

                       

                 

  

               

               

                       

                   

       

       

     

       

     

   

INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

  

INTRODUCTION TO THE 

STATE OF DEEP-SEA 

CORAL AND SPONGE 

ECOSYSTEMS OF THE 

UNITED STATES 

Thomas F. 

Hourigan1*, 

Peter J. 

Etnoyer2, and 

Stephen D. 

Cairns3

  

1 NOAA Deep Sea Coral

Research and Technology

Program, Office of Habitat

Conservation, Silver

Spring, MD  

* Corresponding Author:

Tom.Hourigan@noaa.gov

2 NOAA Center for Coastal

Monitoring and

Assessment, National

Centers for Coastal Ocean

Science, Charleston, SC  

3 National Museum of

Natural History,

Smithsonian Institution,

Washington DC

I. Introduction

Large, long‐lived, sessile organisms contribute structural

complexity to seafloor habitats and play an important role in

marine ecosystems. In deep or cold oceanic waters, corals and

sponges are the most important organisms forming such biogenic

habitats (Roberts et al. 2009, Buhl‐Mortensen et al. 2010, Hogg et al.

2010, Rossi et al. 2017). They increase the physical heterogeneity of

habitat, provide refuge and substrate, increase the number and

availability of micro‐habitats for other organisms, and thereby

create hotspots of biological diversity in the deep sea.

Deep‐sea corals, also known as cold‐water corals, have become a

major focus of new deep‐sea research and conservation, both in the

United States and worldwide. Recent reviews (Hovland 2008,

Roberts et al. 2009, Cordes et al. 2016a) have highlighted the value

of the habitats they create and their vulnerability to anthropogenic

impacts.

In comparison to deep‐sea coral habitats, deep‐sea sponge grounds

have, until recently, been relatively overlooked and poorly

understood (Hogg et al. 2010). This too is beginning to change as

new research has highlighted the extent and importance of these

habitats (Maldonado et al. 2016).

Squat lobster on an

Iridogorgia octocoral in

the Johnston Atoll

component of the Pacific

Remote Islands Marine

National Monument.

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

  

Deep‐sea coral and sponge ecosystems have

become a major focus of international

conservation efforts. United Nations General

Assembly resolutions (UNGA Resolutions

61/105, 64/72, and 66/68) have identified cold‐

water coral habitats as vulnerable marine

ecosystems in need of protection from

significant adverse impacts of deep‐sea bottom

fishing on the high seas. In response,

international guidelines on deep‐sea fishing

(FAO 2009) and conservation actions by

Regional Fishery Management Organizations

worldwide have focused on protecting coral

and sponge habitats as vulnerable marine

ecosystems. The Conference of the Parties to the

Convention on Biological Diversity (Decision

IX/20: CBD 2008) adopted scientific criteria

(Annex I to the decision) for identifying

ecologically or biologically significant marine

areas in need of protection in the open ocean

and deep sea. Deep‐sea coral and sponge

habitats meet the criteria for such designation.

The State of Deep‐Sea Coral and Sponge Ecosystems

of the United States presents new information

gathered over the last decade in the U.S., and

summarizes how this information is

increasingly being used to inform our nation’s

ocean resource management. This introduction

describes the purpose and purview of the

report, and provides a brief summary of

national‐level activities over the last decade that

have supported progress in research,

conservation and management.

II. About This Report

In 2007, the United States National Oceanic and

Atmospheric Administration (NOAA)

published the first peer‐reviewed report on the

State of Deep Coral Ecosystems of the United States

(Lumsden et al. 2007; hereinafter referred to as

the “2007 Report”). The 2007 Report

summarized research on these communities in

U.S. waters up to 2006, focusing on the biology

and importance of structure‐forming deep‐sea

corals and the communities they support, the

threats they face, and their distribution and

conservation status in U.S. waters. In the

decade since 2007, there has been a tremendous

expansion of interest in the science and

management of these ecosystems in the U.S.

and internationally.

The State of Deep‐Sea Coral and Sponge

Ecosystems of the United States serves as an

update to the 2007 Report. It consists of six

regional chapters that cover new information

on research and efforts to conserve deep‐sea

coral ecosystems since 2007. The regional

chapters also provide the first summary of

research on deepwater sponge ecosystems –

though for most regions this information is

relatively limited. This volume does not include

a U.S. Caribbean chapter, as there has been

relatively little new information from waters

surrounding Puerto Rico, the U.S. Virgin

Islands, and Navassa Island since information

from that region was last reviewed (Lutz and

Ginsburg 2007). Each chapter is accompanied

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

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Figure 1. The 2017 State of Deep‐Sea Coral and Sponge Ecosystems of the United States provides an update

to the first NOAA report, State of Deep‐Sea Coral Ecosystems of the United States: 2007.

by an online list of deep‐sea coral species

known from that region. These independently

citable and peer‐reviewed lists (including a U.S.

Caribbean list) update species inventories

contained in the 2007 Report, and substantially

increase the number of taxa recorded in U.S.

waters. NOAA will work with the taxonomists

to update these online deep‐sea coral species

lists regularly and supplement them with

similar species lists for deep‐sea sponges.

The 2007 Report contains a large amount of

background information on regional

ecosystems and management efforts prior to

2007, and the current report is not meant to

replace this. Rather, it builds on the 2007

Report, and provides an update on new

research and management efforts that have

occurred through 2016.

In addition to the regional update chapters, this

report includes six spotlight chapters that

highlight cross‐cutting themes. Each chapter is

written by leading experts with an emphasis on

how research conducted in the U.S. has

contributed globally to our understanding of

deep‐sea coral species discovery (Cairns et al.,

Chapter 2), population connectivity (Morrison

et al., Chapter 12), predictive modeling

(Guinotte et al., Chapter 8), age and growth of

deep‐sea corals (Prouty et al., Chapter 10),

fishing impacts (Rooper et al., Chapter 4), and a

case study on managing black coral harvests

(Wagner et al., Chapter 6).

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

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Box 1. Defining Deep‐Sea Corals

and Sponges

In this report, “deep‐sea corals” and “deep‐sea

sponges” are defined as corals or sponges that

do not depend upon symbiotic algae and light

for their metabolic requirements, and generally

occur at depths below 50 m (NOAA 2010).

Deep‐sea corals are also referred to as cold‐

water corals (e.g., Roberts et al. 2009) and were

called deep corals in the 2007 Report (Lumsden

et al. 2007).

Structure‐forming deep‐sea corals and sponges

are those larger species that provide three‐

dimensional structure above the seafloor that

can be used as habitat by other species. In the

case of corals, these include both deep reef‐

building stony corals (e.g., Lophelia pertusa), as

well as gorgonians, gold corals, and black

corals, which often have branching tree‐like

forms and either occur singly or occur in

aggregations that increase their habitat value.

The most important deep‐sea structure‐forming

sponge species are those in the classes

Demospongiae and Hexactinellida.

III. Corals and Sponges: Key

Components of Deep-Sea

Biogenic Habitats

Deep‐sea corals and sponges occur throughout

the world’s oceans. Many species attain large

sizes and occur in sufficient densities to create

habitat for numerous associated organisms,

thereby forming the basis for remarkably

complex and fragile benthic communities.

These habitat‐forming or structure‐forming

species (NOAA 2010) act as “ecosystem

engineers” (Jones et al. 2007). Rossi et al. (2017)

have dubbed deep‐sea coral and sponge

habitats “marine animal forests,” due to the

structural and functional similarities of these

communities with terrestrial forests. The three‐

dimensional features formed by many deep‐sea

corals and sponges provide habitat for

numerous fish and invertebrate species and

thereby enhance the biological diversity of

many deepwater ecosystems. There is

increasing evidence that these habitats may

play important ecosystem functions, acting as

hotspots of carbon and nutrient recycling in the

food‐limited deep ocean (Cathalot et al. 2015,

Maldonado et al. 2016).

In addition to habitat and ecosystem functions,

deep‐sea corals and sponges are also valuable

to humans in their own right. Cnidarians

(predominantly octocorals) and especially

sponges are the most important sources of

marine natural products (Leal et al. 2012).

Mehbub et al. (2014) reviewed new sponge‐

derived natural products from 2001 to 2010,

which represented about 29% of all marine

natural products discovered during this

decade. Bath sponges have been harvested for

centuries, but now other sponges are being

studied for insights into new industrial

products ranging from fiber optics (Aizenberg

et al. 2005) to nanocrystals (Morse 2007). Black,

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

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pink, and red corals are the basis for a large

jewelry industry (Wagner et al., this volume).

III.1. Deep-Sea Corals

Deep‐sea corals, also referred to as cold‐water

corals, are a taxonomically and morphologically

diverse group of cnidarians distinguished by

their predominant occurrence in deep or cold

oceanic waters. Cairns (2007) defined corals as

“animals in the cnidarian classes Anthozoa and

Hydrozoa that produce either calcium

carbonate (aragonitic or calcitic) secretions

resulting in a continuous skeleton or as

numerous, usually microscopic, individual

sclerites, or that have a black, horn‐like,

proteinaceous axis.” Table 1 shows the major

taxa of deep‐sea corals. The anthozoan

hexacorals include stony corals (Order

Scleractinia), black corals (Order Antipatharia),

and several species parazoanthid gold corals

(Order Zoantharia – in the genera

Kulamanamana [formerly Gerardia)] and Savalia;

Sinniger et al. 2013).

The anthozoan octocorals include the true soft

corals, stoloniferan corals, gorgonians (Order

Alcyonacea), sea pens (Order Pennatulacea),

and helioporids (the shallow‐water blue coral

and the deepwater lithotelestids in the order

Helioporacea). Recent molecular phylogenetic

studies indicate that the anthozoan subclass

Octocorallia is likely monophyletic, but the

orders (Alcyonacea, Pennatulacea, and

Helioporacea) within the octocorals are likely

not (McFadden et al. 2010). Most modern

taxonomists treat the large and morphologically

diverse soft and gorgonian corals as the single

order, Alcyonacea (Daly et al. 2007). Here, as in

the 2007 Report, we continue to treat the

gorgonians (alcyonaceans with a proteinous

and/or calcitic supporting skeletal axis; i.e.,

species currently included in the suborders:

Scleraxonia, Holaxonia, and Calcaxonia)

separately from the other alcyonaceans (true

soft corals and stoloniferans). We do this for

practical reasons (discussed in Hourigan et al.

2007), since many gorgonians are major

structure‐forming species (in contrast to soft‐

bodied alcyonaceans, which typically are not),

and many surveys report corals as

“gorgonians” based on gross morphology when

species or family‐level identifications are

lacking. This practice also allows comparison to

the 2007 Report. In the online species lists,

however, we have included the gorgonians in

the order Alcyonacea in keeping with generally

accepted taxonomic reviews (Fabricius and

Alderslade 2001, Daly et al. 2007, Watling et al.

2011).

Corals in the class Hydrozoa (sometimes called

hydrocorals) are only distantly related to other

corals (in class Anthozoa). Most deep‐sea

species are limited to a single family,

Stylasteridae (the stylasterid or lace corals, in

the order Anthoathecata). The order Hydrozoa

also includes the calcified shallow‐water fire

corals (Family Milleporidae), and three species

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of long horn corals, only one of which lives in

deep water (Cairns 2007).1

1 A few species of other branching deepwater hydrozoans 

produce chitinous skeletons (e.g., Hydrodendron 

gorgonoide, Order Leptothecata), reach large sizes, and 

may provide habitat functions similar to many deep-sea 

corals. While morphologically-similar to gorgonians, 

these are currently not considered to be corals. 

Although more than 600 species of scleractinian

corals occur deeper than 50m (Cairns 2007),

most are solitary corals and only about 20 are

considered framework‐forming (constructional)

species that contribute to deepwater coral reefs

or bioherms (Roberts et al. 2009). The six most

significant species are Lophelia pertusa,

Solenosmilia variabilis, Goniocorella dumosa,

Oculina varicosa, Madrepora oculata, and

Enallopsammia profunda (Roberts et al. 2009). In

U.S. waters, deep‐sea coral bioherms

constructed primarily by O. varicosa, L. pertusa,

and E. profunda occur in the Southeast U.S. and

by L. pertusa in the Gulf of Mexico. Deep‐sea

coral reefs support faunal communities that are

much higher in biomass and diversity than

surrounding unstructured deep‐sea habitats

(Cordes et al. 2008, Roberts et al. 2009, Rossi et

al. 2017). Deepwater reefs may also provide an

important link between the benthos and diel

vertical migrating mesopelagic fishes and

macronekton invertebrates (Gartner et al. 2008,

Davies et al. 2010).

Since 2007, there has been an increased focus on

other types of deep‐sea coral habitats, both in

the U.S. and internationally. This includes high

density aggregations of gorgonians or black

corals, often referred to as coral “gardens,” and

groves of sea pens (Buhl‐Mortensen et al. 2010,

2017; Auster et al. 2013; Stone et al. 2005; Stone

2014; De Clippele et al. 2015; Pérez et al. 2016).

These have much broader depth and

geographic distributions than deep‐sea stony

coral reefs, and have also been recognized as

important biodiversity hotspots in the deep sea

(Buhl‐Mortensen et al. 2017).

III.2. Deep-Sea Sponges

Sponges are sessile animals in the phylum

Porifera, and are among the oldest lineages of

animals (Hooper and van Soest 2002). Most

species are marine, found from tropical to polar

environments and from very shallow to abyssal

depths (van Soest et al. 2012). There are four

extant classes: Demospongiae (the largest class

– sometimes referred to as siliceous and horny

sponges), Homoscleromorpha (recently

separated from the demosponges), Calcarea

(calcareous sponges) and Hexactinellida (glass

sponges) (Table 2). The World Porifera

Database (Van Soest et al. 2017) lists over 9575

extant species of marine sponges (7,742

Demospongiae, 878 Hexactinellida, 834

Calcarea, and 121 Homoscleromorpha). The

phylogeny, systematics, and taxonomy of

sponges have recently undergone extensive

revisions, and many aspects remain unresolved.

The Systema Porifera (Hooper and van Soest

2002) represented a major systematic revision of

the phylum. This revision was supplemented

by recent major revisions to the orders,

Homoscleromorpha (Gazave et al. 2010),

Demospongiae (Morrow and Cárdenas 2015),

and Hexactinellida (Dohrmann et al. 2017).

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

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Table 1. Corals in the phylum Cnidaria that occur in deepwater (> 50 m).

Class Subclass Order

Common

Names

Habitat Contribution

Scleractinia Stony corals

A few branching species form deep‐water

biogenic reef frameworks known as bioherms,

coral banks, or lithoherms. Most deep‐sea

species are small solitary cup corals.

Hexacorallia Antipatharia Black corals

Many branching forms, some of which can

reach large sizes. Often co‐occur with

gorgonians.

Zoantharia Gold corals

Only a few species in the family

Parazoanthidae form rigid skeletons. They

parasitize other corals and need other coral

hosts to settle on. Gold corals can live for over

2000 years.

Anthozoa

Alcyonacea

True soft corals

and stoloniferan

corals

Soft‐bodied species. Most are small and

although they can occur in significant

densities do not appear to be major structure‐

forming species.

Octocorallia

Gorgonacea

(= Alcyonacea,

in part)

Gorgonians

Many branching forms that can reach large

sizes. A number of species can occur in dense

aggregations.

Pennatulacea Sea pens

Unlike most other coral orders, sea pens are

mostly found on soft sediments, where they

can form dense beds that provide important

habitat.

Helioporacea Lithotelestids

Only three species in one genus are known

from deep water. Contribution to habitat is

unknown.

Hydrozoa Hydroidolina Anthoathecata

Stylasterids or

lace corals

(Family

Stylasteridae)

Can form branching colonies. Most species

are relatively small. May be confused with

stony corals but the resemblance is

superficial.

Longhorn

hydrozoans

Only one species (in the Family

Hydractiniidae) of this group is known from

deep water. Not an important structure‐

forming species.

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Deep‐sea sponges can play ecological roles

similar to those of deep‐sea corals, creating

significant three‐dimensional structure on the

sea floor that is used by numerous species

(Freese and Wing 2003, Bell 2008, NOAA 2010,

Buhl‐Mortensen 2010, Stone 2014, Hogg et al.

2010, Maldonado et al. 2016). Although they

can be found on many different bottom types,

most occur on hard substrata (van Soest et al.

2012), also favored by most deep‐sea corals.

Individual sponges can host a rich complement

of microorganisms (Taylor et al. 2007, Webster

et al. 2012) and serve as habitat for a variety of

larger taxa, including both commensal and

obligate symbionts (Klitgaard 1995, Buhl‐

Mortensen 2010). For example, Sedberry et al.

(2004) reported 947 invertebrates representing

ten taxonomic groups living in just five

individual deepwater sponges of different

genera collected in the Southeast U.S. region.

Sponge aggregations can range from small

patches to dense “sponge grounds” in many

deep‐sea areas. These deep‐sea sponge grounds

remain poorly mapped and understood,

prompting Hogg et al. (2010) to christen them

“Cinderellas of the deep seas.”

Demosponge Aggregations: A variety of

demosponges can create monospecific or

multispecies aggregations. In the Aleutian

Islands of Alaska, demosponges greatly

outnumber corals and are a primary component

of highly diverse coral and sponge gardens

(Stone et al. 2011).

Demosponges in the order Tetractinellida

(formerly order Astrophorida) can form dense

and extensive aggregations (commonly known

as astrophorid sponge grounds) on gravel and

coarse sand bottoms from 150 – 1,700 m deep in

cold temperate and arctic regions (Maldonado

et al. 2016). Off Norway, the most abundant

sponges on these grounds (e.g., Geodia barretti)

can reach sizes of 1 m and biomasses as high as

45 kg/m2 (Kutti et al. 2013). Similar sponge

grounds are found along the continental shelf

and slopes off Labrador and Newfoundland

(Murillo et al. 2012; Knudby et al. 2013; Beazley

et al. 2015). Smaller aggregations of

tetractinellid sponges are common in the deep

sea at lower latitudes. Other unique types of

deep‐sea demosponge aggregations include

“lithistid” sponge grounds and carnivorous

sponge grounds (Maldonado et al. 2016).

Glass Sponge Reefs and Aggregations: In the

northeast Pacific off British Columbia, glass

sponges (class Hexactinellida) in the order

Sceptrulophora form unique sponge reefs up to

19 m high and many km long at depths of 90‐

240 m (Conway et al. 2001, 2005). Smaller glass

sponge reefs have recently been documented in

Southeast Alaska (Stone et al. 2014, Stone and

Rooper, this volume). Elsewhere, glass sponges

can form dense, sometimes monospecific,

aggregations principally at depths below 300 m

(Maldonado et al. 2016). In abyssal depths,

small glass sponges are among the few

organisms providing refuge for other species

(Beaulieu 2001).

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Table 2. Sponges in the phylum Porifera that occur in deep water (> 50 m).

Class Subclass Order

Common

Names

Habitat Contribution

Heteroscleromorpha 18 Orders

Demosponges

Demosponges are a large,

diverse group. Many species

reach large sizes and along

with glass sponges represent a

major structure‐forming taxon

in deep water.

Demospongiae Keratosa

Dendroceratida

Dictyoceratida

Verongimorpha

Chondrillida

Chondrosiida

Verongiida

Amphidiscophora Amphidiscosida

Glass sponges

Glass sponges along with

demosponges represent the

primary structure‐forming

deepwater taxa. A few species

form large reefs or bioherms in

Southeast Alaska and British

Columbia.

Hexactinellida

Hexasterophora

Hexasterophora

incertae sedis

Lychniscosida

Lyssacinosida

Sceptrulophora

Calcarea

Calcaronea

Baerida

Leucosolenida

Lithonida Calcareous

sponges

Most calcareous sponges are

found in shallow water. A few

species occur in deeper water.

Calcinea

Clathrinida

Murrayonida

Homoscleromorpha ‐‐ Homosclerophorida ‐‐

A small group of mostly

encrusting forms in deep

water.

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Figure    2. Dense                    community of glass sponges on Pioneer Seamount in the Northwestern    Hawaiian Islands.

INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

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Dense aggregations of filter‐feeding deep‐sea

sponges may also play an important ecosystem

function in nutrient and biogeochemical cycles.

They filter large amounts of water and can

convert dissolved organic matter into

particulate organic matter, which in turn is

used by other organisms (Maldonado et al.

2016). In this way, sponges may play an

important role in carbon, nitrogen, and silicate

cycling and enhancing local productivity.

There is international recognition that deep‐sea

sponge grounds represent vulnerable habitats.

Deep‐sea sponges have been recognized as a

key component of vulnerable marine

ecosystems (FAO 2009) and create habitats that

meet the criteria for Ecologically and

Biologically Significant Areas in the deep sea

(Hogg et al. 2010). Deep‐sea sponge ecosystems

also face many of the same threats as deep‐sea

corals – particularly damage from bottom

trawling (Freese et al. 1999, Freese 2003,

Wassenberg et al. 2002, Hogg 2010, Stone and

Rooper, this volume). Deep‐sea sponge

aggregations are a habitat type listed on the

OSPAR list of Threatened and/or Declining

Species and Habitats (OSPAR 2008). This

recognition led NOAA to include deep‐sea

sponges in its 2010 Strategic Plan for Deep‐Sea

Coral and Sponge Ecosystems (see below and

Box 2).

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11

IV. U.S. National Overview

IV.1. A Strategic Approach

The National Oceanic and Atmospheric

Administration (NOAA) is the lead federal

agency mandated to conserve and manage the

nation’s marine resources, including deep‐sea

coral and sponge ecosystems. In 2010, NOAA

published a Strategic Plan for Deep‐Sea Coral and

Sponge Ecosystems: Research, Management, and

International Cooperation (NOAA 2010, Box 2).

The plan identifies goals, objectives, and

approaches to guide NOAA’s research,

management, and international cooperation

activities on deep‐sea coral and sponge

ecosystems.

Of particular consequence was the Strategic

Plan’s approach to managing bottom‐fishing

impacts to deep‐sea coral and sponge habitats.

Bottom‐contact fishing gears, especially bottom

trawls, currently present the most important

and widespread threat to deep‐sea coral and

sponge habitats, both worldwide (Roberts et al.

2009, Hogg et al. 2010, Ragnarsson et al. 2017)

and within many U.S. regions (Hourigan et al.

2007, Rooper et al., this volume). Because

NOAA’s National Marine Fisheries Service

(NMFS), in partnership with the regional

Fishery Management Councils, is the federal

agency responsible for managing fisheries in

the U.S. exclusive economic zone (EEZ) where

most deep‐sea corals and sponges occur,

managing fishing threats to these ecosystems is

a primary focus of the Strategic Plan.

Box 2. Strategic Plan

NOAA’s 2010 Strategic Plan guides the agency’s

objectives and approaches in three areas related

to deep‐sea coral and sponge ecosystems:

1. Exploration and Research –provides

decision‐makers with scientific information

to enable effective ecosystem‐based

management.

2. Conservation and Management – guides

NOAA efforts to enhance protection of these

ecosystems, working with the Regional

Fishery Management Councils, other Federal

agencies and partners. NOAA’s strategy is

based on authorities provided through the

Magnuson‐Stevens Fishery Conservation

and Management Act (MSA) and the

National Marine Sanctuaries Act.

3. International Cooperation – describes

NOAA’s participation in international

activities to study and conserve vulnerable

deep‐sea coral and sponge ecosystems.
                           

     

           

       

       

             

         

             

           

           

         

             

           

           

                 

         

           

             

         

               

               

             

           

               

                 

               

         

           

         

     

               

       

             

             

           

         

         

             

         

           

             

           

                 

       

         

               

       

       

             

           

             

             

             

         

           

           

               

             

           

       

     

       

             

         

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The NOAA Strategic Plan supports area‐based

(i.e., spatially‐explicit) protection of identified

areas of high density structure‐forming deep‐

sea corals or sponges, and recommends a

precautionary approach to prevent expansion

of the most damaging fishing activities into

unsurveyed areas that might contain deepwater

corals, sponges, and other vulnerable biogenic

habitats (Hourigan 2014). This approach

formed the basis of the historic protection

measures proposed by the Mid‐Atlantic Fishery

Management Council and instituted by NOAA

in 2016 (see Packer et al., this volume). The

Strategic Plan also highlighted the importance

of measuring and addressing fisheries bycatch

of deep‐sea corals and sponges. The NMFS

National Bycatch Reduction Strategy (NMFS

2016a) calls upon the agency to: (1) identify

areas of high bycatch of deep‐sea corals and

sponges; (2) to work with regional Fishery

Management Councils and the fishing industry

to close these areas to high‐bycatch gears as

called for in the Strategic Plan for Deep‐Sea Coral

and Sponge Ecosystems; and (3) to collect better

data on coral bycatch and post‐interaction

mortality. The agency’s most recent U.S.

National Bycatch Report (NMFS 2016b)

contains quantitative information on the

bycatch of deep‐sea corals and sponges off the

West Coast and Alaska.

Within the U.S. government, interest in these

deepwater ecosystems is not limited to NOAA.

The U.S. Geological Survey (USGS) released

Strategic Science for Coral Ecosystems 2007‐2011

(USGS 2007), which described the information

needs of resource managers for both shallow

and deep coral ecosystems and summarized

research

partners. The agency‐shared long‐term vision is

to develop a more complete understanding of

the physical, chemical, and biological processes

– both natural and anthropogenic – that control

or influence the structure, function, and

ecological relationships within coral

communities.

conducted by USGS scientists and

IV.2. Research Advances

Understanding

IV.2.i – Deep-sea science spurred by

advances in technology

Research on U.S. deep‐sea coral and sponge

ecosystems has benefited from the availability

of new tools and techniques (Fig. 3). In 2008,

NOAA commissioned the Okeanos Explorer to

systematically explore our largely unknown

ocean for the purpose of discovery and the

advancement of knowledge. Telepresence uses

satellite communications to allow scientists

from around the world to participate in

expeditions remotely by connecting the ship

and its discoveries live with audiences ashore.

The NOAA Ship Okeanos Explorer is joined by

the Ocean Exploration Trust’s E/V Nautilus, also

equipped with telepresence capabilities, and a

new generation of NOAA fisheries research

vessels conducting deep‐sea coral and sponge

research in U.S. waters. These and other vessels

have begun to map the seafloor more

systematically, and at higher resolution, using

multibeam sonar. Meanwhile, improvements to

remotely‐operated vehicles (ROVs),

autonomous underwater vehicles (AUVs) and

other equipment (Fig. 3) have provided for

more detailed surveys, revealing previously

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  

Figure 3. Advanced Technology used to explore and understand deep‐sea coral and sponge ecosystems: a) NOAA

Ship Okeanos Explorer; b) Deep Discoverer ROV; c) Telepresence‐enabled exploration; d) SeaBed AUV.

unknown habitats to depths of 6000 m.  

In addition to new survey technologies,

understanding of these ecosystems has

benefitted from a host of other new approaches.

As reviewed by Cairns et al. (this volume), new

genetic techniques have revolutionized our

understanding of taxonomy and systematics,

and are being applied for the first time to

understanding the connectivity of deep‐sea

coral and sponge populations (e.g., Morrison et

al., this volume). Predictive modeling of deep‐

sea coral habitats has advanced considerably,

and is helping target both new research and

conservation efforts (Guinotte et al., this

volume). Other new techniques allow corals to

tell the history of past oceanographic conditions

they have experienced (Prouty et al., this

volume), and reveal the remarkable microbial

associates of deep‐sea corals (e.g., Kellogg et al.

2016) and sponges. Sponges, in particular, host

exceptionally dense and diverse microbial

communities (reviewed by Taylor et al. 2007,

Webster et al. 2012, Thomas et al. 2016).

IV.2.ii – NOAA’s Deep Sea Coral Research

and Technology Program

NOAA’s Deep Sea Coral Research and

Technology Program is the only U.S. national

program dedicated to research on deep‐sea

coral ecosystems. It was established in the 2007

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  

Box 3. National Database for Deep‐Sea Corals and Sponges

(https://deepseacoraldata.noaa.gov)

NOAA’s Deep Sea Coral Research and Technology Program has compiled a database of the known locations of deep‐sea

corals and sponges, beginning in U.S. waters (Hourigan et al. 2015). Representing the most comprehensive collection of

deep‐sea coral and sponge records and information for U.S. waters, the database is available publicly in NOAA’s Deep‐

Sea Coral Data Portal. The portal includes a digital map displaying more than 500,000 records. The National Database

includes records from samples archived in museums and research institutions, reported in the scientific literature, as

well as observations collected during deep‐water surveys conducted by NOAA and other research institutions.

In addition to showing locations of corals and sponges, the fully searchable map also provides access to the following:

 In situ photos of the organisms.

 Extensive associated data available for download about coral and sponge observations, including record

provenance, details about where and how they were observed or collected, and, where available, ecologically

important information, such as their density, size, and habitat.

 Reports that characterize the deep‐sea coral and sponge habitats surveyed over the past decade by scientists

from NOAA, other agencies, and universities.

 Deep‐sea coral habitat suitability model layers.

The National Database for Deep‐Sea Corals and Sponges is continually expanding, incorporating new records from

recent fieldwork observations and historic archives quarterly. Additional software tools for data exploration and

analysis are under development. The Portal also offers information about studies funded by the Deep Sea Coral

Research and Technology Program since 2009 and a growing library of NOAA publications on deep‐sea corals and

sponges.

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

  

reauthorization of the Magnuson‐Stevens

Fishery Conservation and Management Act

(MSA, Section 408), the nation’s primary

fisheries management legislation. The mission

of the program is to provide the science needed

to conserve and manage vulnerable deepwater

ecosystems. While focused on deep‐sea corals,

NOAA has informed congress and the public

that the program will also collect

complementary information, if available, on

high biodiversity deep‐sea sponge habitats

(NMFS 2008).

The Deep Sea Coral Research and Technology

Program began operations in 2009. It has

conducted 3‐4 year field research initiatives in

nearly all U.S. regions, as outlined in the

regional chapters that follow. Surveys

conducted by the program and its partners

have supported management efforts across the

country, including identification of vulnerable

coral and sponge habitats to be protected from

damaging bottom‐fishing gears, expansion of

National Marine Sanctuaries and Monuments,

and the establishment of the Northeast

Canyons and Seamounts Marine National

Monument.

These field initiatives have been supplemented

by targeted projects to map deep‐sea coral

distributions, model predicted deep‐sea coral

habitat (Guinotte et al., this volume), study

coral genetics and connectivity (e.g., Everett et

al. 2016), and support coral bycatch reduction.

Data collected by the Deep Sea Coral Research

and Technology Program and its partners

contribute to NOAA’s National Database for

Deep‐Sea Corals and Sponges (Box 3, Hourigan

et al. 2015) and are available through the

program’s map portal

(www.deepseacoraldata.noaa.gov).

IV.2.iii – Other major research programs

The last decade also saw the results from major

U.S. interagency collaborations focused on

deep‐sea coral ecosystems led by the

Department of Interior’s Bureau of Ocean

Energy Management (BOEM, formerly

Minerals Management Service), in collaboration

with the U.S. Geological Survey (USGS) and

NOAA, and sponsored by the National

Oceanographic Partnership Program (NOPP).

As described by Boland et al. (this volume),

major multidisciplinary studies were conducted

in the Gulf of Mexico during the 2004‐2006

(Lophelia I; Sulak et al. 2008), 2005‐2009 (Chemo

III; Brooks et al. 2014) and 2008‐2012 (Lophelia II;

Brooks et al. 2016). These studies, particularly

Lophelia II, produced unprecedented new

information on the biology and life history of

major structure‐forming corals (Lophelia pertusa,

the black coral Leiopathes sp., and the gorgonian

Callogorgia spp.), community structure, trophic

relationships, and other aspects of these deep‐

sea coral communities (Brooks et al. 2016). A

similar collaboration was conducted in the Mid‐

Atlantic’s Baltimore and Norfolk Canyons

(Packer et al., this volume).

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

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IV.3. Conservation Status of U.S.

Deep-Sea Coral and Sponge

Ecosystems

Understanding the conservation status of deep‐

sea coral and sponge ecosystems requires

information on the following topics: (1) the

spatial distribution of these biogenic habitats;

(2) the spatial extent and intensity of

anthropogenic activities that pose potential

threats and their overlap with biogenic habitats;

(3) the sensitivity of these ecosystems to

different impacts and their recovery potential;

and (4) the effectiveness of management

measures to address these threats (Ragnarsson

et al. 2016). There has been substantial progress

over the last ten years on each of these fronts in

regard to deep‐sea coral and sponge

ecosystems.

IV.3.i – Spatial distribution of U.S. deep-sea

corals and sponges

Deep‐sea habitats are difficult and expensive to

survey. The United States has the world’s

second largest exclusive economic zone (EEZ),

most of it below the edge of the continental

shelf (i.e., greater than ~200 m deep). This area

remains largely unmapped, and the areas

visually surveyed for deep‐sea corals or

sponges are miniscule. Nevertheless, the last

decade has seen a more systematic approach to

both mapping the seafloor and understanding

the distribution of deep‐sea habitats. This

information has been identified as the first

priority nationally for management (Hourigan

2014, regional chapters in this Report).

Deep‐Sea Coral Distributions: Structure‐forming

corals are widespread in deeper waters of all

regions except the U.S. Arctic. Although largely

unexplored, only sea pens and one soft coral

(Gersemia sp.) have been reported from the

Chukchi Sea and Beaufort Seas (Stone and

Rooper, this volume). This result contrasts to

the Arctic north of the Atlantic, where extensive

and relatively diverse coral habitats have been

discovered off Canada, Greenland, and Norway

(Roberts et al. 2009). NOAA’s National

Database for Deep‐Sea Corals and Sponges

(Box 3) has resulted in the first comprehensive

maps of coral presence in areas of U.S. waters

that have been sampled (map annexes in each

regional chapter). Predictive habitat models

allow some extrapolation of these data to

unsurveyed areas (Guinotte et al., this volume).

Such maps and models of coral presence,

however, do not yet capture the local extent of

habitats nor the density and diversity of corals

within the habitats – features that are most

important for determining their conservation

value.

Although deep‐sea corals occur widely, areas of

high‐density aggregations (e.g., coral

“gardens”) are highly localized, and may be

small (many on the scale of tens to hundreds of

meters across). They therefore represent a

comparatively rare habitat type. Yet these coral

garden areas support diverse communities of

other organisms and represent hotspots of

biological diversity in the deep sea (e.g., Auster

et al. 2013, Stone 2014). Their diversity and

rarity makes them both extremely valuable and

extremely vulnerable.

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

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The only true deepwater stony coral reefs

(bioherms) have been observed in the U.S.

Southeast (Hourigan et al., this volume) and

Gulf of Mexico (Boland et al., this volume), and

most recently on seamounts of the

Northwestern Hawaiian Islands and Emperor

Seamount Chain (Baco et al. 2017). Some deep‐

sea reef formations may also occur in

unsurveyed areas of the U.S. Caribbean. U.S.

deepwater coral reefs are most diverse and

numerous in the Southeast, where they

probably rank among the most extensive deep‐

sea coral reef provinces in the world. These

reefs, along with coral and sponge gardens in

the Aleutian Islands, represent the largest

extents of highly diverse U.S. deep‐sea coral

communities. However, every U.S. region

contains truly remarkable habitats, often in

areas of clear water – on ridges, seamounts,

canyon walls and shelf‐edge breaks – where

there is hard substratum, sufficient food, and

moderate to strong currents.

New explorations continue to reveal amazing

new habitats. Deep‐sea surveys in the U.S.

Pacific Island Territories began in 2016 (too

recent to be reflected in the U.S. Pacific Islands

summary; Parrish et al., this volume), revealing

extensive and dense coral and sponge gardens

on ridges and seamounts. New research has

also uncovered unexpectedly rich habitat areas

within the current bottom‐fishing footprint that

appear to have escaped damage (e.g., coral

gardens in the Gulf of Maine, Northern

California, and Aleutian Islands). These areas

represent conservation priorities, as they face

the most immediate threats from bottom‐

fishing.

Deep‐Sea Sponge Distributions: The Gulf of

Mexico (Rützler et al. 2009) and the Aleutian

Islands in Alaska (Stone et al. 2011) are the only

regions with moderately systematic lists of

deep‐sea sponge species, although species lists

exist for certain sub‐areas (e.g., California, Lee

et al. 2007). Mapping of sponge distributions

has barely begun, and with the exception of

some Alaskan areas (Rooper et al. 2014, Rooper

et al. 2016), no predictive habitat models have

been developed to date. The most

comprehensive picture of sponge presence

comes from scientific trawl surveys conducted

off Alaska and the U.S. West Coast (Clarke et

al., this volume) – though most records are only

recorded as “Porifera,” and the surveys cannot

access areas of rough topography that may be

especially important habitats for many species.

These surveys indicate that certain areas have

high bycatch of sponges and likely represent

high‐density sponge grounds. These include

monospecific sponge grounds in Alaska’s

Bristol Bay (B. Stone pers. comm.), and highly

diverse sponge gardens in the Aleutian Islands

that have also been visually surveyed (Stone

2014, Goddard et al. 2017). Bycatch of sponges

from commercial fisheries off the U.S. West

Coast and Alaska is an order of magnitude

larger by weight than the bycatch of corals, and

the Alaska sponge bycatch is 50 to 100 times

higher than off the West Coast. This trend

supports the generalization that high‐density

deep‐sea sponge grounds are more common in

cold temperate waters (Maldonado et al. 2016).

Pile and Young (2006) reported that the deep‐

sea glass sponge, Sericolophus hawaiicus, forms

dense beds (mean density: 4.7/m2) over

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

  

Figure 4. Massive glass sponge discovered at a

depth of 2117 m in the Northwestern Hawaiian

Islands. The picture shows the sponge and the

ROV Deep Discoverer. The sponge was estimated

to be over 3.5 m in length, 2.0 m in width and 1.5

m in height, making it the largest sponge recorded

to date.

extensive areas at depths between 360 – 460 m

off the Big Island of Hawaii. Unlike many deep‐

sea sponges that occur only on hard substrata,

S. hawaiicus is adapted to anchor in the sand.

Recent explorations in both the Hawaiian

Archipelago and other U.S. Pacific Islands and

seamounts have revealed dense aggregations of

glass sponges on rocky ridges. These

explorations included the discovery of what

may be the largest sponge ever reported

(Wagner and Kelley 2016; Fig. 4).

There have been no systematic surveys of deep‐

sea sponge habitats in other U.S. regions. Our

understanding of these ecosystems has been

hampered by lack of appreciation of their

importance, and by limited U.S. expertise in

taxonomy and ecology of deepwater sponges.  

IV.3.ii – Anthropogenic threats

The 2007 Report summarized information on

anthropogenic threats to deep‐sea coral

ecosystems in U.S. regions (Hourigan et al.

2007). Bottom trawl fisheries were the most

serious threat in Alaska, the U.S. West Coast,

Northeast, and Southeast regions. Other

bottom‐tending gear, including traps, bottom‐

set longlines, and gillnets can also damage

deep‐sea corals (Baer et al. 2010, Sampaio et al.

2012, Rooper et al., this volume). These gears

may be used preferentially in steep and rocky

habitats (i.e., areas of high rugosity) that are

inaccessible for trawling, thereby representing

the primary fishing gear damaging corals and

sponges in such areas. Oil and gas development

was considered a moderate threat in the Gulf of

Mexico, and invasive species and precious coral

harvests were of particular concern in Hawaii.

Other threats, while possibly significant at a

local level, had relatively small footprints

compared to bottom fishing. At the time, there

was insufficient information on potential

impacts of climate change to these ecosystems

to assign a threat level.

The last decade has seen an increase in

awareness of potential threats to deep‐sea

ecosystems (Ramirez‐Llodra et al. 2011,

Mengerink et al. 2014, Koslow et al. 2016).

Ramirez‐Llodra et al. (2011) concluded that

impacts to the deep sea were increasing

globally, with deep‐sea coral habitats among

the most vulnerable, and fishing, especially

bottom trawling, being their most serious

current threat. Climate‐related changes,

including ocean acidification, ocean warming,

and changes in deep‐sea current regimes and

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

  

productivity were expected to become major

threats in the future. This general conclusion

based on scientific expert opinion is supported

by recent reviews of threats to deep‐sea coral

ecosystems by other authors (Roberts et al.

2009, Cordes et al. 2016a, Koslow et al. 2016,

Ragnarsson et al. 2017), each of which

highlighted vulnerability to threats from

fishing, fossil fuel exploitation, climate change,

and ocean acidification. Reviews of impacts to

deep‐sea sponge ecosystems have also

identified bottom‐trawling as the most serious

current impact (Hogg et al. 2010, Maldonado et

al. 2016).

Table 3 provides an updated summary of

anthropogenic threats to deep‐sea corals and

sponges in U.S. regions based on published

literature and expert judgement (reviewed in

the regional Chapters), and compares these to

threats described in the 2007 Report. We

assume that impacts to sponges from physical

disturbances are qualitatively similar to impacts

to corals from the same activities (e.g., Stone

2014). The following represent the major

changes to the 2007 threat assessment:

Bottom Fishing: Damage from bottom trawling is

still considered the biggest threat to deep‐sea

coral and sponge ecosystems where it occurs in

U.S. regions where these gears are used

(Alaska, U.S. West Coast, Northeast U.S.).

Bottom trawling in the Southeast U.S. and Gulf

of Mexico is restricted to a small number of

vessels engaged in deepwater shrimp fisheries.

There is still incomplete information on the

footprint of bottom‐fisheries in the U.S., but

information has improved in certain areas (e.g.,

the West Coast; see Clarke et al., this volume).

In a series of National Bycatch Reports (NMFS

2011, 2013, 2016b), NOAA quantified the

bycatch of corals and sponges by fishery in

Alaska (2003‐2005; 2010‐2013) and the U.S. West

Coast (2011‐2013). There continues to be

significant bycatch of corals and sponges –

primarily from a limited number of trawl

fisheries and from relatively discrete locations

within these large regions. The highest rates by

far are from the rockfish trawl fishery in the

Aleutian Islands. As noted by Rooper et al. (this

volume), fixed gears (e.g., bottom‐set longlines,

gillnets, and traps) can also damage deep‐sea

corals and sponges, but less is known about the

extent of their impacts. Their footprint is

certainly orders of magnitude smaller than that

of trawling, but may allow targeting of prime

coral or sponge habitats that are unsuitable for

trawling. Steps taken by the South Atlantic and

Mid‐Atlantic Fishery Management Councils

have significantly increased the area of

protected deep‐sea coral and sponge habitats,

reducing the threat from bottom‐fishing

impacts to the most important areas.

Oil and Gas Development: The potential impacts

of oil and gas development came into stark

focus with the Deepwater Horizon oil spill

(Boland et al., this volume). Deep‐sea coral

habitats at three sites from 6‐22 km away from

the wellhead (White et al. 2012, Fisher et al.

2014a, Fisher et al. 2014b) were damaged by the

oil spill – evidently as a result of a deepwater

plume. Gorgonians at mesophotic depths (60–

88 m) in areas below the surface oil slick also

exhibited significant declines in condition

19
INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED ST

  

Table 3. Summary of perceived levels of threats (based on Lumsden et al. 2007 and chapters within this report) to deep‐sea 

(2007 and 2017) and sponge communities (2017) for U.S. regions. NA = Not Applicable (i.e., this threat is prohibited or do

within the region).Note: Perceived threat levels reflect only the occurrence of these stressors in a region and their potential, if unmitigated, to 

and sponge communities. They do not indicate the actual impacts of each stressor, which can vary widely within and amon

location of deep‐sea coral and sponge habitats is incompletely known, there is uncertainty over their degree of overlap with 

U.S. has taken substantial management steps to mitigate many threats, and the change in perceived threats for fishing in th

Southeast reflect recent protections. The 2007 Report did not separate ocean acidification from climate change, and deep‐se

analyzed as a potential threat.

U.S. REGIONS

THREATS Alaska West Coast Pacific Islands Northeast Southeast Gulf of Mexi

2007 2017 2007 2017 2007 2017 2007 2017 2007 2017 2007 2

Bottom Trawl

Fishing Impacts High High High High NA NA High Medium High Medium Low ‐ Medium Lo

Me

Other Bottom

Fishing Impacts

Low ‐

Medium Medium Low ‐ Medium Low ‐ Medium Low Low Low ‐ Medium Low ‐ Medium Low ‐ Medium Low ‐ Medium Low ‐ Medium Lo

Me

Deep‐Sea Coral

Harvest

NA NA NA NA Medium Medium NA NA NA NA NA N

Oil & Gas

Development Low Low Low Low NA NA NA Low NA Low Medium Me

Cable Deployment Low Low Low Low Unknown Low Low Low Low Low Low L

Sand and Gravel

Mining Low Low NA NA NA NA Low Low Low Low Low L

Deep‐Sea Mining Low Low Low ‐ Medium Low Low L

Invasive Species Unknown Unknown Unknown Unknown Medium Medium Unknown Unknown Unknown Unknown Unknown Unk

Climate Change

Unknown

Unknown

Unknown

Unknown

Unknown

Unknown

Unknown

Unknown

Unknown

Unknown

Unknown

Unk

Ocean Acidification Medium Medium Medium Low Low ‐ Medium Lo

Me

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(Etnoyer et al. 2016, Silva et al. 2016). Cordes et

al. (2016b) recently reviewed the environmental

impacts of the oil and gas industry: when

potential accidental spills are taken into

account, the potential threat posed by oil and

gas development in the Gulf of Mexico to deep‐

sea coral ecosystems is greater than originally

supposed by Hourigan et al. (2007). The

Deepwater Horizon oil spill was a rare, worst‐

case scenario, and greatly improved measures

have been put in place to prevent this kind of

spill from happening again. During the last

decade, offshore oil and gas exploration and

leasing has been considered for additional

regions, including Alaska (Stone and Rooper,

this volume) and the Mid‐Atlantic (Packer et

al., this volume). While active fossil fuel

development in these regions is currently on

hold, the increased possibility of this moving

forward in the future is reflected in Table 3.

Renewable Energy: Offshore renewable energy,

especially offshore wind energy, has recently

become a major driver for more comprehensive

ocean planning in a number of regions. The

nation’s first offshore wind installation began

operations off Rhode Island in 2016 (Packer et

al., this volume). Most offshore wind facilities

are expected to be sited on the continental shelf

relatively close to shore, and thus are less likely

to impact major deep‐sea coral and sponge

habitats. There is, however, the potential for

anchored wind turbines in deepwater areas

nearshore (e.g., in Hawaii), which could affect

deep‐sea coral and sponge habitats. Developers

have also proposed potential marine current

energy off southeastern Florida (Vinick et al.

2012) and ocean thermal energy conversion

projects off Hawaii and southeastern Florida

that could damage deepwater biogenic habitats.

Any proposed activities would result in site‐

specific surveys conducted to avoid impacts

from installations.

Deep‐Sea Mining: There are currently no

proposals for deep‐sea mining within U.S.

waters. Nevertheless, there is increasing

interest and capacity for deep‐sea mining

worldwide. Deep‐sea mineral resources contain

commercially important quantities of high‐

grade ores increasingly valued in modern

technology (Hein 2010, Hein et al. 2013). The

principal deep‐sea mineral resources being

considered for mining include the following:

 Polymetallic manganese nodules, generally

occurring at abyssal depths (3,500 – 6,000

m).

 Seafloor massive sulfides, also known as

polymetallic sulfides, associated with active

or extinct hydrothermal vents.

 Cobalt manganese crusts on seamounts. The

prime crustal zone occurs in the North

Pacific, including areas around Hawaii and

U.S. Pacific territories.

 Phosphorite nodules, typically found

between 200‐400 m depth.

Mining, if it occurs, is likely to completely

destroy deep‐sea coral or sponge habitats

within its footprint (Ramirez‐Llodra et al. 2011,

Levin et al. 2016). Additional impacts are

expected from sediment plumes produced

during mining operations. Currently the

greatest concern in U.S. waters appears to

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

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impacts from mining to the particularly rich

deep‐sea coral and sponge habitats on

seamounts in the U.S. Pacific Islands (Parrish et

al., this volume). Many of these occur in the

Prime Crust Zone of the Central Pacific

(Schlacher et al. 2014), which contains large

concentrations of commercially valuable deep‐

sea minerals (Hein et al. 2013).

Climate Change and Ocean Acidification: There are

still many unknowns concerning the potential

impacts of climate change and ocean

acidification on deep‐sea coral ecosystems. The

2007 Report did not assign a level of threat to

these ecosystems in the United States from

climate change due to a lack of information

(Hourigan et al. 2007). Since then, the Fifth

Assessment Report of the Intergovernmental

Panel on Climate Change (IPCC) concluded

that ocean warming has affected deep‐sea

ecosystems at least down to 2000 m (Hoegh‐

Guldberg et al. 2014). Effects of warming on

deep‐sea coral and sponge communities include

direct impacts on survival (e.g., Brooke et al.

2013, [coral]) and growth (Stone et al. 2017,

[coral]), and an array of indirect effects linked

to increasing water temperature (Sweetman et

al. 2017). These include decreased dissolved

oxygen concentrations (Keeling et al. 2010,

Levin and Le Bris 2015), altered hydrodynamics

(Birchenough et al. 2015), or decreased

productivity of surface waters and export of

food to the deep‐sea (Jones et al. 2014).

Rising atmospheric carbon dioxide (CO2) is also

directly responsible for ocean acidification with

associated changes in carbonate chemistry that

affect coral calcification (Guinotte et al. 2006).

Ocean acidification results from net uptake by

the ocean of CO2 emissions, which decreases

carbonate ion concentration in ocean waters

(Feely et al. 2004). Worldwide, ocean waters

from 200–3000 m are expected to face the

largest reductions in pH by the year 2100

(Sweetman et al. 2017). This decrease has been

forecast to hamper production of biogenic

carbonates (aragonite and calcite) in the

skeletons of corals, and is likely to be most

problematic for reef‐forming stony corals. The

deep waters of the northeast Pacific have the

shallowest aragonite and calcite saturation

horizon, and may provide a unique

opportunity to study the response of deep‐sea

corals to ocean acidification. Corals in this

region could be surviving in potentially

corrosive water conditions for some months of

the year (Feely et al. 2008).

In Table 3 we have divided climate impacts into

two categories – climate changes and ocean

acidification. We identify the latter as a low to

medium threat to deep‐sea corals currently, but

is likely to become a high threat in the future.

Threats from ocean acidification are highest

where the aragonite saturation horizon is

shallowest (in the North Pacific) and where

there are deep‐sea scleractinian coral reefs,

which are expected to be especially vulnerable

to dissolution. These factors come together in

the newly discovered deepwater reefs in the

Northwestern Hawaiian Islands and Emperor

Seamounts (Baco et al. 2017). There are likely

synergies between ocean acidification and other

impacts of climate change.

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Compared to corals, there is little information

on potential responses of sponges to climate

change and ocean acidification. Kahn et al.

(2012) observed changes in abyssal sponge

populations correlated with climate‐driven

changes in particulate organic carbon. Bennett

et al. (2017) presented initial experiments with

shallow‐water sponges that found that high

temperatures may adversely affect sponge

survival. In their experiments, elevated partial

pressure of carbon dioxide (pCO2) alone did not

result in adverse effects, however it exacerbated

temperature stress in heterotrophic sponges,

but not in phototrophic species. We are not

aware of similar experiments with deep‐sea

sponges, all of which are heterotrophic.

IV.3.iii – Vulnerability of Deep-Sea Coral

and Sponge Ecosystems to Threats

Most deep‐sea corals and sponges are highly

vulnerable to physical impacts such as those

from bottom trawling (Roberts 2009, Hogg

2010). Recovery from damage will depend on

extent of the damage, and the ability of the

damaged organisms to recover or for new

recruits to settle and grow. This, in turn, is

governed by the life‐history characteristics of

deep‐sea corals and sponges, which tend to

include slow growth, late maturity, extreme

longevity, and infrequent recruitment events,

all of which make these organisms particularly

slow to recover from disturbances (Roberts et

al. 2009, Hogg et al. 2010).

Since the 2007 Report there has been growing

evidence for extreme age and slow growth of

deep‐sea corals (Prouty et al., this volume).

Roark et al. (2009) reported that Hawaiian gold

corals and one species of black coral could

reach ages of 2,500 to over 4,000 years,

respectively. Even relatively shallow‐water

holaxonian corals in Alaska may take 60 years

to reach maximum size (Stone et al. 2017).

There have been few studies of recruitment of

deep‐sea corals in U.S. waters, but most

evidence points to relatively low and episodic

recruitment (Cordes et al. 2016a).

Clark et al. (2016) found that corals and

sponges were highly vulnerable to fishing

impacts, and their life history attributes meant

that, once damaged, the recovery potential of

biogenic habitats was highly limited. They

concluded that recovery would take decades to

centuries after fishing had ceased. There have

been few studies investigating recovery of

damaged deep‐sea coral habitats in U.S. waters,

but observations to date indicate that its

potential is very limited (Stone and Rooper, this

volume).

Sponges are also extremely vulnerable to

damage from fishing gears (Freese et al. 1999,

Freese 2001, Stone 2014, Stone et al. 2014,

Maldonado et al. 2016, Malecha and Heifetz

2017). Suspended sediments associated with

bottom trawling may also adversely affect

deep‐sea sponges (Tjensvoll et al. 2013). Less is

known about the life history of sponges than of

deep‐sea corals, however, there is growing

evidence that many species are slow‐growing

and long‐lived. The massive shallow‐water

demosponge, Xestospongia muta from the

Florida Keys, is estimated to live for more than

2300 years (McMurray et al. 2008). The deep‐sea

glass sponge, Monorhaphis chuni, may hold the

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

  

distinction of being the longest‐lived animal on

earth. Jochum et al. (2012) measured oxygen

isotopic ratios and Mg/Ca ratios from a giant

basal spicule of a specimen collected at 1110 m,

and estimated that the sponge had been

growing for 11,000 ± 3,000 years.

The glass sponge, Aphrocallistes vastus, is widely

distributed in the northern Pacific Ocean from

Panama through the Bering Sea and to Japan

(Stone et al. 2011). Austin et al. (2007)

concluded that juveniles of this species can

reach a moderate size within 10‐20 years on

glass sponge reefs off British Columbia, but

may require a century to reach full size. They

also found that that the species was very

susceptible to physical damage and that

recruitment appeared to be rare at their study

site. Kahn et al. (2016) observed recruitment,

growth, and response to damage of glass

sponge reefs over a three year period. They

observed recruitment, as well as growth of

sponges ranging from 0‐9 cm/year, but sponges

did not recover from experiments simulating

larger scale damage. The authors concluded

that the sponge reefs are not resilient to

disturbances such as bottom trawling.

In conclusion, research over the last decade has

provided increased evidence for the high

vulnerability and low resilience of deep‐sea

coral and sponge habitats and the communities

they support. This conclusion emphasizes the

value of management measures to protect

remaining undamaged deep‐sea coral and

sponge communities.

IV.3.iv – Actions to conserve U.S. deepsea coral and sponge ecosystems

The 2007 Report summarized steps taken

within U.S. waters to manage impacts to deep‐

sea corals and other deepwater habitats. These

efforts primarily focused on NOAA’s National

Marine Sanctuaries and area‐based fishing gear

closures developed by the regional Fishery

Management Councils. The latter addressed the

most immediate threat to vulnerable benthic

habitats: a few relatively small areas were

specifically established to protect deep‐sea coral

habitats, e.g., the Oculina Banks Habitat Area of

Particular Concern (established in 1983 and

expanded in 2000), Alaska Sitka Pinnacles

Marine Reserve (2000), and Aleutian Islands

and Gulf of Alaska Coral Habitat Conservation

Areas (2006). Additional deepwater areas in

U.S. waters have been established for general

habitat protection (Sutter et al. 2013).

New Areas Protected from Fishing: The 2007

Magnuson‐Stevens Act reauthorization

recognized the importance of deep‐sea coral

habitats and provided new discretionary

authority to protect these habitats in their own

right (MSA Sec. 303(b)(2)). Since then, Fishery

Management Councils in each region have

actively included deep‐sea coral ecosystems in

discussions of conservation measures

(Hourigan 2014). Large‐scale, area‐based

conservation measures have specifically

targeted deep‐sea coral ecosystems for

protection. In 2010, the South Atlantic Fishery

Management Council established five

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

  

deepwater Coral Habitat Areas of Particular

Concern that protect deepwater coral reefs in

an area of 62,717 km2 (Hourigan et al., this

volume). The Mid‐Atlantic Fishery

Management Council was the first to use the

new MSA discretionary authority to protect

deep‐sea coral habitat regardless of formal

recognition as Essential Fish Habitat (Packer et

al., this volume). The Council proposed the

designation of the Frank R. Lautenberg Deep‐

Sea Coral Protection Area, encompassing more

than 99,000 km2 (~38,000 square miles) in 2015.

NMFS approved this amendment in 2016,

establishing the largest protected area in the

U.S. Atlantic. This conservation approach was

based on NOAA’s Strategic Plan for Deep‐Sea

Coral and Sponge Ecosystems, protecting specific

canyons where deep‐sea corals had been found,

as well as freezing the footprint of most bottom

fisheries to prevent expansion into new

deepwater habitats. The New England Fishery

Management Council is exploring major deep‐

sea coral protections in 2017 using the same

approach and authority.

New and Expanded National Monuments and

Sanctuaries: New discoveries of rich deep‐sea

coral and sponge habitats have also resulted in

other important advances in deep‐sea

conservation. In the U.S. Pacific Islands,

Presidential proclamations 8335‐8337 (January

2009) designated three new National

Monuments: the Marianas Trench Marine

National Monument, Rose Atoll Marine

National Monument, and Pacific Remote

Islands Marine National Monument. The Pacific

Remote Islands Marine National Monument

and the existing Papahānaumokuākea National

Marine Monument (established in 2006) were

subsequently expanded by Presidential

proclamation in 2014 and 2016, respectively.

These new and expanded Monuments protect

important deep‐sea and seamount habitats and

form the largest network of marine protected

areas in the U.S. (Parrish et al., this volume). In

2016, the first marine national monument in the

Atlantic Ocean, the Northeast Canyons and

Seamounts Marine National Monument, was

established under the authority of the

Antiquities Act of 1906 (Packer et al., this

volume). This new monument protects several

submarine canyons and the four New England

seamounts in the U.S. EEZ. The Presidential

Proclamation specifically referenced deep‐sea

corals, along with “other structure‐forming

fauna such as sponges and anemones,” as

resources that “create a foundation for vibrant

deep‐sea ecosystems” and are extremely

sensitive to disturbance from extractive

activities.

Several of NOAA’s national marine sanctuaries

have also undergone major expansions in the

last decade. In 2008, NOAA incorporated

Davidson Seamount — a volcanic seamount

that is home to rich deep‐sea coral and sponge

habitats — into the Monterey Bay National

Marine Sanctuary, providing comprehensive

management in addition to the 2006 EFH

bottom‐gear closure. In 2012, the Fagatele Bay

sanctuary expanded to protect five additional

areas and became the National Marine

Sanctuary of American Samoa. In 2015, NOAA

expanded the boundaries of Cordell Bank and

Gulf of the Farallones National Marine

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

  

Sanctuaries to an area north and west of their

old boundaries, to include new deepwater areas

surveyed by the Deep Sea Coral Research and

Technology Program. These expansions were

motivated, in part, by the discovery of

important deep‐sea coral habitats. Most of the

existing sanctuaries have also been actively

exploring the deeper extents of their protected

areas (e.g., Clarke et al., this volume, and

Boland et al., this volume) and incorporating

results into their management plans.

Offshore Energy Management: The Bureau of

Ocean Energy Management (BOEM) manages

renewable energy development in federal

waters. The Deepwater Horizon disaster has

prompted review and strengthening of offshore

oil and gas regulations (Boland et al., this

volume). Mesophotic and deep‐sea coral

habitats will also be a priority for restoration

activities in the Gulf of Mexico over the coming

decade under the Deepwater Horizon Oil Spill

Final Programmatic Damage Assessment and

Restoration Plan. The last decade has also seen

the first offshore leases for wind energy

development off the U.S. East Coast (Packer et

al., this volume). While these offshore wind

farms do not extend into deep water, this

development has become a major driver for

regional ocean management planning efforts. A

number of regions have begun to incorporate

deep‐sea coral observation and predicted

habitat information into their broader plans to

protect vulnerable ecosystems (e.g., the Mid‐

Atlantic Regional Council for the Ocean).

In summary, the last decade has seen deep‐sea

biogenic habitats, especially deep‐sea coral

habitats, taking an increasingly central role in

ocean planning and conservation in every U.S.

region. Interest in these ecosystems seems

likely to continue as we learn more about their

value and vulnerability.

V. Conclusions

Deep‐sea coral research over the decade since

the 2007 Report on the State of Deep Coral

Ecosystems of the United States has become more

targeted, systematic, and collaborative. As

described in the chapters that follow, this trend

has resulted in tremendous advances in our

understanding of the distribution of many taxa,

as well as insights into the basic biology and

ecology of these organisms. In contrast,

knowledge of deep‐sea sponges remains

rudimentary at best, despite our increasing

recognition of their importance to deep‐sea

ecosystems. Even a basic understanding of the

life history of the most important structure‐

forming species and their distribution in U.S.

waters continues to elude researchers and

managers.

The new research has led to increased

awareness of the beauty, ecological importance,

and fragility of deep‐sea ecosystems. This

awareness is manifested in new conservation

measures directed toward the deep sea,

especially deep‐sea coral habitats. Most

important among these have been the new

Marine National Monuments in the U.S. Insular

Pacific and Northeast U.S., and large new

fishery management zones that will protect

over 175,000 km2 of deep‐sea areas off the U.S.

East Coast, including many important deep‐sea

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INTRODUCTION TO THE STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE UNITED STATES

  

coral habitats. Deep‐sea sponge grounds have

received no specific protections, though many

are likely included in these recent large‐scale

conservation areas that address bottom‐fishing.

As marine research and management move

forward, conservation of these remarkable

ecosystems will be enhanced by continued

mapping of deep‐sea biogenic habitats, and

research focused on understanding their

structural diversity, ecological function, and

contribution to biodiversity and ecosystem

productivity. Each of the following chapters

contains recommendations for future research.

The next steps in conservation will use this

improved understanding to apply a more

targeted approach to identifying high priority

areas for protection. Based on the success in

“freezing the footprint” of the most damaging

fishing gears, future progress will likely require

management within existing fishing areas,

using a collaborative approach with fishers and

other resource users that promotes sustainable

use while protecting the most valuable benthic

communities. Future progress will also need to

address increasing economic uses of deep‐sea

resources (e.g., deep‐sea mining) and the

potential impacts of a changing climate.

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Everett MV, Park LK, Berntson EA, Elz AE,

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Newly described species of stylasterid

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2015 on a CAPSTONE expedition to

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Courtesy of the NOAA Office of

Ocean Exploration and Research.
SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •

39

SPECIES DISCOVERY 

OF DEEP-WATER 

CORALS AND 

SPONGES IN U.S. 

WATERS (2007-2016)

I. Discovery of Deep-Water Corals

In order to understand biodiversity and to conserve a biota, 

or to simply converse about it in a meaningful manner, one 

must know the names of the units being discussed – the 

species. The 2007 “State of Deep Coral Ecosystems” report 

(Lumsden et al. 2007) concentrated on the most ecologically 

important species - the structure-forming deep-water corals 

(highlighted in Appendices 1.1, 1.2, and elsewhere). In seven 

articles, each on a different region of the United States (U.S.), 

the authors dutifully included detailed lists of the deepwater coral species found within each region, sometimes 

subdividing their regions into smaller subunits. Yet, there 

was never a comprehensive synthesis or grand tally of how 

many deep-water coral species occur in U.S. waters in the 

2007 report. 

Admittedly, a ‘comprehensive synthesis’ requires some 

time and effort, but this can initially be accomplished by 

comparing the seven previously published regional lists 

(Lumsden et al. 2007) with the new online regional species 

lists associated with this report (Online Annexes). Taking 

into account revisions in taxonomy and eliminating duplicate 

entries, i.e., those species occurring in two of more of the 

seven regions, resulted in a comprehensive list (Online 

Annex)

1

. Table 1 summarizes the number of species in each 

of the five groups of corals known to inhabit the seven 

geographic regions. 

1

 Pacific Islands region is composed of the Hawaiian Islands, Guam, American Samoa, Northern Mariana Islands, and 

Pacific Remote Islands (Baker I., Howland I., Jarvis I., Johnston Atoll, Kingman Reef, Palmyra Atoll, and Wake I.); and 

the Caribbean region is composed only of Puerto Rico, the U.S. Virgin Islands, Navassa I., and Sarranilla Bank.

Stephen D. 

Cairns1*, 

Robert P. 

Stone2

, Ewann 

A. Berntson3,

and Shirley A.

Pomponi4

• • •

1 National Museum of Natural History, Smithsonian 

Institution, Washington, DC

* Corresponding Author:

CairnsS@si.edu

2 NOAA Auke Bay Laboratories, Alaska Fisheries 

Science Center, Juneau, AK

3 NOAA Conservation 

Biology Division, Northwest 

Fisheries Science Center, 

Port Orchard, WA

4 Marine Biomedical & 

Biotechnology Research 

Program, Harbor Branch 

Oceanographic Institute, 

Florida Atlantic University, 

Fort Pierce, FL
40

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •

Table 1. Comparison of numbers of deep-water coral species by region between 2007 and 2016. Gross 

totals include species that overlap in two or more regions. Corrected totals eliminate the overlapping 

species. Taxa not identified to species level not included. 

Figure 1. The number of deep coral species known from four taxonomic groups occurring in US waters as 

of 2007 (left bar, dark blue) and newly identified species as of 2016 (right bar, light blue).

Antipatharia Scleractinia Zoanthidea Octocorallia Stylasteridae

2007 2016 2007 2016 2007 2016 2007 2016 2007 2016

Northeast U.S. 0 1 16 17 0 0 37 47 0 0

Southeast U.S. 2 14 57 77 0 0 42 92 7 15

Gulf of Mexico 17 28 47 72 0 0 44 118 8 9

U.S. Caribbean 9 13 42 45 0 0 23 43 14 14

U.S. West Coast 3 7 17 19 0 0 39 80 4 6

Alaska 10 8 8 9 0 0 45 58 12 23

Hawaii 20 21 53 54 0 1 93 104 4 4

Gross Totals 61 92 237 293 0 1 323 542 49 71

Corrected Totals 45 60 151 163 0 1 247 385 42 53

New taxa since 2007 15 12 1 138 11
41

Among the 662 deep-water corals 

presently known from U.S. waters (Table 

1, and comprehensive species list herein), 

Octocorallia have by far the highest species 

richness (58.2% of the grand total), followed 

by the azooxanthellate Scleractinia (24.6%), 

Antipatharia (9.1%), and the Stylasteridae 

(8.0%). The rank order of these four major 

groups is consistent with three other 

comprehensive lists of deep-water corals 

made for other oceanic regions, specifically the 

Gulf of Mexico (Cairns and Fautin 2009: 354 

species), New Zealand (Cairns et al. 2009: 376 

species), and European waters (Costello et al. 

2001: 261 species). From this table one can also 

calculate that the most species-rich regions are 

the Gulf of Mexico (227 species), southeastern 

US (198 species), and Hawaii (184 species); 

the most depauperate regions being the 

northeastern U.S. (65 species) and Alaska (98 

species). Table 1 can then form the basis for 

what progress has been made between mid2007 to 2016 in describing deep-water coral

species richness of the seven U.S. regions.

Progress is inevitably a function of research

effort. In the U.S., deep-water coral research

is supported by the National Oceanic and

Atmospheric Administration (NOAA), the

Smithsonian, the U.S. Geological Survey

(USGS), the Bureau of Ocean Energy

Management (BOEM), the National Science

Foundation (NSF), academia, and private

industry, among others. These groups are all

invested in the exploration, sampling, and

systematics of deep-water corals and their

habitats. Examples of new species discoveries

are shown in Figures 2a, 2b, and 3.

A regional increase in species richness can

occur in three ways: (1) by the description of

new species, (2) by the discovery of a range

extension into U.S. waters of a previously

described species, and (3) by the addition of

species previously overlooked. It is relatively

Figure 2a. Iridogorgia magnispiralis at 2200 m depth, a new record for the northern Gulf of Mexico, 

from NOAA’s Okeanos Explorer expedition of 2012.

Figure 2b. Isidella tentaculum at 891 m depth, a new species in the Northeast Pacific described in 2008, 

from NOAA’s Gulf of Alaska Seamount Expeditions aboard RV Atlantis in 2002 and 2004.

Figure 3. An in-situ image of a zoanthid gold coral from Hawaii called Kulamanamana haumeaae

(Sinniger et al. 2013). The colony represents a new genus and species of gold coral, previously thought to 

be in a different genus, Salvia or Gerardia. Image credit: Public Library of Science One.

2a 2b 3

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
42

easy to compile a list of the new species for 

a region but somewhat harder to ascertain if 

there has been a range extension, as evidence 

of the latter is often buried in the literature. 

After a thorough review of the scientific 

literature, we have determined that 62 new 

species and three new genera of deep-water 

corals had been newly described from U.S. 

waters between 2007 and 2016. In addition, 

numerous species had been reported as a 

result of range extensions, and as having been 

previously overlooked. The rate of 62 new 

species in ten years equates to 6.2 new species 

per year, or an annual worldwide increase in 

described species of 0.19% (6.2/3336 known 

deep-water coral species; Cairns 2007b). This 

is considerably lower than the annual increase 

in new species descriptions for Scleractinia 

over a 30-year period, as calculated by Cairns 

(1999) as 1.23%, although his rate applied to 

both deep- and shallow-water species around 

the world, and the 0.19% rate is limited to 

the deep-water species from U.S. waters. 

Table 1 and Figure 1 show that the most 

actively growing group was the Octocorallia, 

increasing by 138 names, or 55.9%, since 2007.

The relatively high number of new octocorals 

is due, at least in part, to the large size of the 

group, their poorly known taxonomy, and 

to the revisionary papers on the fauna of: (1) 

the New England Seamounts (Watling 2007, 

Simpson and Watling 2011, Watling and 

France 2011, Pante and Watling 2011, Cairns 

2007c); (2) the Hawaiian Islands (Cairns and 

Bayer 2007, Cairns 2009, 2010); (3) the U.S. 

west coast (Cairns 2007c); and (4) the Alaskan 

waters (Cairns 2011, Cairns and Baco 2007). 

Most of those specimens have been collected 

by NOAA research vessels. Also, many new 

records of all taxonomic groups were based on 

as yet unpublished museum records and other 

gray literature reports. A revisionary work 

by Cairns and Lindner (2011) on the Alaskan 

stylasterids served to increase the number 

of species in this deep-water coral group by

26.2%. Relatively few species were added

to the U.S. antipatharian and scleractinian

fauna, their components increasing by 33.3%

and 7.9%, respectively. It is of interest to note

that of the 62 new species described during

this period, 16 (26%), were additionally

characterized by molecular sequencing,

usually of mitochondrial genes (Cairns and

Baco 2007, Etnoyer 2008, Watling and France

2011, Pante and Watling 2011, Cairns and

Lindner 2011, Opresko et al. 2012, and Tu et al.

2016).

Three faunistic lists were also published

during this period, which helped to increase

the number of species in U.S. waters: (1)

Felder and Camp (2009), all taxa at all depths

for the Gulf of Mexico; (2) DeVictor and

Morton (2010), octocorals from the South

Atlantic Bight from 0-200 m; and (3) Watling

et al. (2011), octocorals at all depths and

regions, the latter allowing for cross-checking

of taxonomic names of the Atlantic regions, a

guide to some of the range extensions for these

regions, and the addition of some species

that had been overlooked in the 2007 report.

Based on Watling et al. (2011) eight species

were added to the U.S. fauna (5 octocorals and

3 stylasterids), mainly from the southeastern

region. The total number of deep-water corals

now known from U.S. waters is 662 species,

up 177 (36.5%) from the 485 species listed in

2007 (Table 1), representing 19.8% of the total

known deep-water coral fauna (Cairns, 2007b).

Sixty-two of the 177 new names were newly

described (see the comprehensive species list)

and 115 had been previously overlooked.

In summary, we believe that a significant

amount of work is being done to discover

new species, document range extensions, and

to uncover previously overlooked records of

deep-water corals in U.S. waters leading to

numerous descriptions, especially in the group

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
43

Octocorallia. We expect that future increases 

in species descriptions will likely fall in that 

group. We also acknowledge that it is not the 

lack of collections that hinders advancement 

but the lack of manpower that limits progress 

and causes uneven growth (Cairns 2007b). A 

pertinent example of this is the dissertation by 

K. Muzik (1979), containing the description

of 18 new species of Hawaiian deep-water

octocorals, which remains unpublished.

II. Species Discovery of Deep- as 

Ad

Water Sponges pr

Sponges are one of the simplest, oldest-living 

metazoan phyla and one of the most abundant 

groups of benthic marine organisms (Fig. 

4). They are more diverse than corals, and 

single- or multi-species sponge aggregations 

may play a similarly important role in the 

ecology of deep-sea ecosystems (Maldonado et 

al. 2016). At present, there are approximately 

8500 described species with perhaps as 

many yet to be discovered. Worldwide, 

about 50 new species are described each 

year (Cárdenas et al. 2012). The systematics 

of sponges are particularly challenging 

given their ancient origin, simple body 

organization, complex microbial associations 

and morphological plasticity. As a result, 

phylogenetic relationships within the Porifera 

remain mostly unresolved, and this hampers 

progress in basic studies of sponge biology 

and biodiversity, including comparative 

evolutionary studies that use sponge species 

as model organisms as well as efforts to 

conserve or economically exploit sponges. 

The deep-water sponge fauna of the U.S. has 

not been inventoried as systematically as the 

deep-water coral fauna. There are important 

historical collections dating back decades 

or more for most regions but there have 

been few dedicated efforts in recent years

to collect sponges beyond those provided

opportunistically. The 2006 Magnuson-Stevens

Reauthorization Act established the Deep Sea

Coral Research and Technology Program, but

made no specific provision to consider deepwater sponges, and they were not included in

the 2007 NOAA Report (Lumsden et al. 2007).

Still, deep-water sponges are abundant in the

same depth range as deep-water corals, have

complex structure that provides habitat for

other fauna, and face many of the same threats

deep-water corals (e.g. Hogg et al. 2010). 

ditionally, sponges and their associates 

oduce a diverse array of chemicals that are 

of interest for biotechnology (Pomponi et al. 

2007). Accordingly, the NOAA Strategic Plan 

for Deep-sea Coral and Sponge Ecosystems 

(2010) provides nearly equal emphasis on the 

deep-sea sponge and coral fauna, and clear 

guidance to include sponges in research and 

management efforts. 

Whereas sponges have taken a back seat 

to their more charismatic deep-water 

counterparts, there is strong evidence that 

poriferans provide important habitat for fish 

and invertebrates. With that in mind, sponges 

were specifically collected during submersible 

cruises in the Aleutian Islands of Alaska in 

2003–2004. This was done, in part, to construct 

a field guide (Stone et al. 2011) and to help 

establish a useful fisheries bycatch monitoring 

program. Ninety-four (94) sponge species 

were collected during this period, and 28 

(30%) were new to science! This high rate of 

new species discovery, and many new range 

extensions, clearly indicated that the sponge 

fauna of the region is poorly documented. In 

response to these findings, NOAA scientists 

boarded regional groundfish survey vessels 

during the summer of 2012 and, with the 

new guide in hand, collected 118 unfamiliar 

bycatch specimens. These new collections 

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
44

will undoubtedly yield many new discoveries 

and provide insight into the true species 

diversity of the region. Since 2004, more casual 

collections in other regions of Alaska (e.g., the 

Bering Sea and Gulf of Alaska) have yielded 

additional species that have either been 

recently described or are awaiting 

formal description.

More than 55 taxa, principally hexactinellids, 

have been opportunistically collected along 

the Pacific Coast as part of NOAA’s Deep 

Sea Coral Research and Technology Program 

three-year research initiative in that region 

(Clarke et al., this report). Approximately 

28 taxa (51%) are believed to be new to 

science and await formal identification. 

Thirty hexactinellid specimens were recently 

collected during an expedition to the Necker 

Ridge in the Northwest Hawaiian Islands 

and 19 taxa, or 72%, are believed to be new to 

science. Other collections in the Pacific Islands 

Region (principally funded by NOAA through 

the Hawaii Undersea Research Laboratory) 

included 10 new species, all believed to be 

new to science (Chris Kelley, University of 

Hawaii at Manoa, personal communication). 

Together, these regional collections represent 

a new species discovery rate of about 60%. 

This rate could be somewhat inflated since 

researchers may preferentially collect forms 

that are unfamiliar to them. Regardless, these 

data support those from Alaska and clearly 

indicate that the deep-water sponge fauna of 

the Pacific Ocean is poorly documented and 

that a large percentage of the total fauna are 

yet to be discovered.

One particularly noteworthy example of 

unexplored and undocumented deepwater 

sponge fauna was the exciting discovery of 

the carnivorous harp sponge Chondrocladia 

lyra off the coast of California (Lee et al. 2012) 

(Figure 6). This unusual group of sponges, to 

date found only in the Class Demospongiae, 

Family Cladorhizidae, differs drastically in 

body plan and cellular organization from 

the rest of the Phylum Porifera, and was 

only recently discovered (Vacelet and BouryEsnault 1995). Since that time, at least 133 (as 

of 2014) new species of carnivorous sponges 

have been described, primarily in deep 

water (Cristobo et al. 2015).Using MBARI 

ROVs Tiburon, Ventana and Doc Ricketts, four 

Figure 4. Sponge gardens 

(areas of particularly 

diverse and abundant 

sponge fauna) were 

discovered in the central 

Aleutian Islands in 

2004. Seven species of 

sponges are visible in this 

photograph, including 

two new to science and 

recently described.

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
45

Figure 5. Undescribed species of Geodia from 118 meters depth off the coast of Florida, collected by 

HBOI-FAU using the Johnson-Sea-Link submersibles in 2007.

new species were collected off the west coast 

of the U.S. (Lundsten et al. 2014) since 2007, 

increasing the total of U.S. Pacific carnivorous 

sponge species from 7 to 11 and several 

more are in the process of being described 

(Lundsten, MBARI, pers. comm.).

Many sponge collections have been made 

during the past decade in the South Atlantic 

Bight and Gulf of Mexico with funding 

from NOAA. Harbor Branch Oceanographic 

Institute-Florida Atlantic University has 

collected and archived more than 1200 deepwater sponges from U.S. waters using the 

Johnson-Sea-Link manned submersibles (Reed 

et al. 2005, 2006). Rützler et al. (2009) identified 

339 sponge species in a checklist of the Gulf 

of Mexico, of which 66 (48 demosponges and 

18 hexactinellids) were identified as deep-sea 

sponges. Species lists have been submitted 

to NOAA as part of the technical (cruise) 

reports and several new species from these 

collections await formal identification (Fig.5). 

A new record of occurrence of the carnivorous 

sponge Chondrocladia verticillata was 

discovered in 2011 off the Florida Keys at a 

depth of 850 m by the Cooperative Institute for 

Ocean Exploration, Research, and Technology 

at HBOI-FAU, using the University of 

Connecticut ROV Kraken. Recent interest in 

resolving the phylogenetic relationships of the 

phylum Porifera has resulted in more detailed 

studies of these collections using molecular 

and morphological systematics (Dohrmann et 

al. 2012a, b; Hill et al. 2012). The increasing use 

of molecular techniques is facilitating species 

identification and discovery (Wörheide et al. 

2007, Erpenbeck et al. 2015), and leading to 

significant revisions of sponge systematics 

(e.g., Morrow and Cardenas 2015).

There is a renewed interest in sponge 

taxonomy and systematics resulting from 

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
46

discoveries that continue to be made during 

exploration of poorly studied areas in U.S. 

waters. The discovery of sponge-associated 

microbial symbionts and natural products 

with pharmaceutical potential and the 

application of molecular systematics to resolve 

phylogenetic relationships have added to this 

interest. As with other deep-water fauna, there 

are already long queues in the laboratories 

of the few sponge taxonomists worldwide, 

suggesting that future collections should be 

carefully planned and coordinated with them 

if possible.

III. Genetic Identification of

Deep-water Coral Species

An understanding of the population 

structure of deep-water coral species is 

critical to ascertaining gene flow within 

and among distant populations, in order 

to gauge the effects of habitat loss, and 

meet the directives of the 2006 MagnusonStevens Act and the National Marine 

Fisheries Service (NMFS) federal mandate. 

A species inventory of deep-water corals 

off U.S. coastlines is a necessary first step 

toward a comprehensive understanding 

of the ecology and distribution of this rich 

and diverse faunal assemblage. Fisheries 

bycatch data and benthic trawl surveys can 

help quantify species diversity and outline 

their biogeographical distributions, but such 

estimates require accurate and defensible 

species-level identifications; however, the 

taxonomic identification of deep-water corals 

to the species level can be problematic for 

both morphological and practical reasons 

(see McFadden et al. 2011); in some cases, 

species of octocorals that appear similar 

Figure 6. The carnivorous harp sponge Chondrocladia lyra with 5 vanes was collected with the ROV 

Doc Ricketts off California. This unusual group of sponges was discovered only a few decades ago and 

differs in body plan and cellular organization from all other sponges. © 2009 MBARI.

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
47

may only be distinguished from one another 

morphologically using scanning electron 

microscope (SEM) images of sclerites, 

minute skeletal structures embedded in the 

fleshy tissue of these corals. Some species 

of deep-water corals may be defined from 

a single known type specimen. In other 

cases, the known evolutionary relationships 

of some coral families are so inadequate 

that accurate species-level designations are 

currently impossible. Additionally, many 

deep-water coral specimens are collected in 

trawls or dredges, which may damage the 

specimen or yield partial colonies at best. 

These difficulties, combined with a scarcity 

of trained morphological taxonomists, 

make the expanded development of genetic 

techniques to aid in species identification a 

critical necessity. It is encouraging to see, and 

worth noting, that students are being trained 

in coral taxonomy in federal (e.g., S. Cairns, 

P. Etnoyer) and academic laboratories (e.g., S.

France, P. López-González, and L. Watling).

Nevertheless, the use of genetic information,

when combined with morphology, is currently 

the best approach to deciphering evolutionary 

patterns in these species. 

Voucher collections are currently being 

compiled at NOAA’s Northwest Fisheries 

Science Center (Seattle, WA) and NOAA’s 

Center for Coastal Environmental Health and 

Biomolecular Research (Charleston, SC) to aid 

in a more rapid and accurate identification 

of coral specimens. Such collections include 

one or more specimens of each potential 

species, with taxonomic identifications to 

be confirmed by relevant experts. As part of 

this process, SEM images may be produced 

for the vouchers as necessary, to either allow 

the direct identification of some individuals 

or assist morphological taxonomists with 

their examinations and characterizations of 

unknown specimens. Genomic sequences 

are then generated for each voucher so that 

DNA from additional specimens can be used 

to ascertain their identity in the absence of 

labor intensive SEM imaging or the large 

sample sizes required for some morphological 

Figure7a. An unidentified sponge in the genus Aphrocallistes at 255 m depth on Kidney Bank in 

Southern California. Many unique taxa in the Pacific Region await formal description. Image credit: 

NOAA Southwest Fisheries Science Center, Advanced Survey Technologies Group, La Jolla CA.

Figure 7b. The recently described sponge Latrunculia austini occurs at depths between 30 - 384 m in 

the Northeast Pacific from the Olympic Coast off Washington State to the Gulf of Alaska. These sponges 

have unique compounds that offer biomedical potential to curing disease. Image credit: Bob Stone, NOAA 

Alaska Fisheries Science Center.

7a 7b

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
48

analyses. Choosing which area of a coral’s 

genome to investigate is, however, not 

immediately straightforward.

The selection of an appropriate genetic region 

or regions to uniquely identify individual 

species is critical, requiring a fast enough 

mutation rate to yield sufficient differences 

between species, but not so many differences 

that sequence alignment is difficult. Animals 

contain two types of genetic material in 

their cells, which have both similar and 

differing characteristics: nuclear DNA and 

mitochondrial DNA. In diploid organisms, 

nuclear DNA (nDNA) is inherited from 

both parents—one copy of each gene being 

contributed by each parent—and is present as 

a single set of chromosomes in each somatic 

cell. Mitochondrial DNA (mtDNA) is inherited 

from an organism’s mother. Each cell also 

contains multiple copies mtDNA because 

there are numerous mitochondria whereas 

there is only a single nucleus. Mitochondrial 

genomes tend to evolve more quickly than 

nuclear genomes because, at least in higher 

vertebrates, mitochondria lack the ability to 

“repair” accidental mutations in their DNA 

that arise during cell division. This higher 

rate of evolution is useful when attempting to 

identify genetic regions—termed “markers”—

that exhibit sufficient diversity to distinguish 

individuals on a species level. As a result 

of these attributes, mtDNA sequencing has 

become the standard genetic tool for many 

species identifications. Recent reports, 

however, cite inherent problems with using 

mtDNA to investigate recent molecular 

evolution, and this must be considered when 

choosing an appropriate barcoding region (see 

Bazin et al. 2006, Galtier et al. 2009). Presently, 

the most concerted effort to generate a 

database of species-specific genetic sequences, 

i.e., genetic barcodes, is coordinated by the

Barcode of Life (BOL) initiative

(www.barcodeoflife.org). The stated purpose

for the BOL is to assist in situations where a

specimen may be damaged, thereby making

morphological identification difficult, or in

cases where expert taxonomists are required

to make a definitive identification. Barcoding

would potentially solve the problem of species

identification in both instances, as short DNA

sequences can be generated from minimal

amounts of tissue and used by non taxonomic

experts to determine species designations.

The genetic region targeted for most animal

species is a 648 basepair region of the

mitochondrial cytochrome c oxidase 1 gene

(COI); however, this region is not sufficiently

informative for species identification in

octocorals because this subclass experiences a

slower rate of evolution in their mtDNA than

vertebrates (France and Hoover 2002). One

reason cited for this stability is the presence

of the mitochondrial MutS homolog, msh1

(mtMSH) gene, which has high similarity and

homology to mismatch-repair genes found

in bacterial genomes as well as eukaryotic

nuclear genes (Culligan et al. 2000). If the

MutS homolog is functional in octocorals, that

could explain why the mitochondrial genomes

of octocorals are often not sufficiently variable

to distinguish among species.

Interestingly, the mtMSH gene itself

appears to be highly variable and provides

an alternative to the COI gene for species

identification purposes (France and Hoover

2002). The mtMSH gene has been shown to

be specific for many, but not all, coral species

investigated (France and Hoover 2001). The

addition of other gene regions, such as the

“Folmer region” of COI and an adjacent

intergenic area, have demonstrated good

potential as a substitute to the COI when used

in combination with msh1 at the species level

(McFadden et al. 2011). Despite this increased

resolution, there remain limitations in

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
49

identification of certain deep-water coral taxa. 

Further investigations into using alternative 

genes must continue.

Examples of both the utility and limitations 

of combined genetic and morphological 

identifications are now appearing in the 

literature (e.g., Cairns and Baco 2007; Lapian 

et al. 2007; Wagner et al. 2010; Baco and 

Cairns 2012; Pante et al. 2012). Development 

of the Northwest Fisheries Science Center 

voucher collection exemplifies their utility; 

a number of octocorals frequently collected 

in trawl surveys in the Northeast Pacific 

are initially identified as Swiftia sp. (Family 

Plexauridae) based on overall similarities 

in appearance. These samples may also be 

assigned a shipboard species designation 

within the genus Swiftia based on general 

colony formation and color. Figure 4 depicts 

a group of such individuals, all of which 

were originally identified aboard trawling 

vessels as Swiftia sp., S. simplex, or S. pacifica. 

Identification by a taxonomist through a 

detailed morphological examination including 

SEM images of formations and arrangements 

of sclerites, revealed some of these individuals 

actually represent two additional genera plus 

one additional species of Swiftia, despite their 

superficial similarities (Fig. 8). A neighborjoining tree of pairwise differences of the 

mtDNA sequence data (including the Folmer 

region of COI plus the adjacent intergenic 

region, plus msh1) for these same specimens 

corroborated the taxonomic identities of these 

individuals (Fig. 8). On this phylogenetic 

tree, the horizontal branch lengths indicate 

the relative differences measured between 

individuals. Note that individuals of the 

same species group together in “clusters” 

and that the horizontal branch lengths among 

individuals within the same species are very 

short or even zero (e.g., the Swiftia simplex

and Swiftia spauldingi clusters). Slightly 

longer branch lengths are found between 

species within the same genus (e.g., the

difference between Swiftia simplex and Swiftia

pacifica), and the longest branch lengths are

between different genera (e.g., Leptogorgia

and Eugorgia). Additional specimens from

this group can now be sequenced for the

above genetic regions and compared to the

vouchered sequences on this tree, yielding

taxonomic identifications without requiring

the consultation of a taxonomist for every

new individual.

The use of molecular genetics in species

identification alleviates pressure on the

few, overworked coral taxonomists and

can illuminate species distinctions when

morphological distinctions are absent (as in

the case of cryptic species). As previously

mentioned, approximately 25% of newly

described deep-water coral species included

a molecular characterization of primarily

mitochondrial markers. Further coordination

of morphological taxonomists with genetic

labs could readily contribute molecular

information for the other 75% of newly

described taxa. Many genetics labs across the

country are now capable of generating and

analyzing DNA sequence data, as is evidenced

by the recent explosion in the number of

institutions contributing to the BOL project.

In addition to barcode sequencing, recent

advances in molecular genetic technologies

are currently in use, or being developed, that

may prove invaluable to the investigation of

various aspects of deep-water coral species

identification and genetics. Next-generation

sequencing produces gigabases of data

in mere days, enabling the assemblage

and interrogation of entire genomes in a

comparatively short amount of time. The

generation of such massive amounts of

molecular data also allows for the continued

screening of additional genetic regions

that might have utility for species-level

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
50

Figure 8. Phylogenetic comparison of the Fulmer region of COI plus adjacent intergenic region, plus 

msh1, of putative Swiftia spp. with other Plexauridae from the Northeast Pacific. The dendrogram shows 

three clusters of Swiftia specimens corresponding to three different species. Swiftia pacifica is more 

closely related to S. simplex than S. spauldingi. Adelogorgia phyllosclera is more closely related to 

Swiftia spp. than to two other genera - Leptogorgia and Eugorgia.

SPECIES DISCOVERY OF DEEP WATER CORALS IN U.S. WATERS (2007-2016)

• • •
51

identification as well as population-level 

analyses. Other molecular genetic techniques 

that have relevant application in the genetic 

analysis of deep-water corals include, but 

are not limited to: (1) targeted amplicon 

sequencing (Bybee et al. 2011), which can 

use next-generation sequencing techniques 

(e.g., Roche 454, Illumina, and/or Ion Torrent 

platforms) and associated bioinformatics to 

generate large numbers of DNA barcodes 

simultaneously; (2) environmental DNA 

(eDNA), a relatively new technique that 

can identify the presence of species from 

ambient water samples, reducing the need 

to sample individual corals directly and 

increasing overall sampling opportunities; (3) 

microsatellites (msats), also known as simple 

short repeats (SSR), which are tandemly 

repeated stretches of 2-6 dinucleotides, and 

can be used to measure neutral variation 

among populations as well as help gauge 

the proportion of genetic variation among 

different groups within a single species; 

and (4) RAD sequencing (restriction site 

associated DNA sequencing), a genome 

reduction technique that is often employed to 

identify genetic markers—Single Nucleotide 

Polymorphisms (SNPs) and/or msats—that 

have the potential to distinguish among 

individuals, species, or even populations. 

As we learn more about the genetic 

relationships among deep-water corals, we 

stand to gain a richer understanding of species 

diversity as well as ascertain the severity 

of pressing environmental disturbances 

on the health and vulnerability of habitatforming anthozoans, a benefit that will guide 

the development of regulatory policies to 

protect these remote, productive, and fragile 

ecosystems.

IV. Acknowledgments

This manuscript was made more complete and 

more consistent in style by the reviews of Les 

Watling, Chris Kelley, Geoff Cook, Peter 

Etnoyer, Tom Hourigan, and Debe Tighe.

Online Annex - Comprehensive List of 

Deep-Sea Corals in the U.S. EEZ link: 

https://deepseacoraldata.noaa.gov/

library/2017-state-of-deep-sea-coralsreport 

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      ‐

Red tree coral with a school of juvenile

Pacific cod. Courtesy of the Deepwater

Exploration of Glacier Bay National

Park expedition and UCONN NURTEC.
                     

     

                     

                   

                 

                   

             

                       

                       

                     

                 

                   

                       

               

                   

                 

               

               

           

               

                     

               

                   

                   

                   

                   

   

           

   

     

       

       

  

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Robert P. 

Stone1* and 

Chris N. 

Rooper2

  

1 NOAA Auke Bay

Laboratories, Alaska

Fisheries Science Center,

Juneau, AK

* Corresponding author:

Bob.Stone@noaa.gov

2 Resource Assessment

and Conservation

Engineering Division,

Alaska Fisheries Science

Center, Seattle, WA

STATE OF DEEP-SEA 

CORAL AND SPONGE 

ECOSYSTEMS OF THE 

ALASKA REGION 

I. Introduction

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. The

Aleutian Islands have the highest diversity of deep corals in Alaska,

and possibly in the northern North Pacific Ocean, including

representatives of all major taxonomic groups and at least 50

species or subspecies that may be endemic to that region (Stone and

Shotwell 2007). Additionally, the Aleutian Islands harbor 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).

The Aleutian Islands also support the most abundant deep‐water,

ahermatypic (non‐reef building) coral resources in high‐latitude

systems reported anywhere in the world (Stone 2014).

Stone and Shotwell 2007 (hereafter referred to as the 2007 Report)

highlighted diverse coral communities in Alaskan waters including

141 unique coral taxa with gorgonians and stylasterid corals being

the most species rich and abundant and demonstrated that each

region of Alaska has important and unique coral habitats. Detailed

descriptions of each major coral group were presented along with

Redbanded rockfish

(Sebastes babcocki)

taking refuge in a large red

tree coral (Primnoa

pacifica) detached from

the seafloor in Dixon

Entrance, eastern Gulf of

Alaska.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

figures for each sub‐region based on catch

records from National Oceanic and

Atmospheric Administration (NOAA) research

trawl surveys (1975–2004) and the 2004

sablefish longline survey, some published

records from archived specimens, and in situ

observations (R. Stone, unpublished records).

The process revealed that very few records

existed for the Alaskan Arctic (Chukchi and

Beaufort Seas) and a limited but growing body

of records existed for Gulf of Alaska seamounts.

Both commercial and non‐commercial fish and

invertebrate species had been documented in

deep‐sea coral habitat at varying spatial

degrees, but most documented associations

were suspected to be facultative rather than

obligatory.

The 2007 Report also concluded that the major

stressor on deep coral communities in Alaska

was commercial fishing activities with bottom

trawls exhibiting the highest threat based on

severity of effects, extent of effects, geographic

extent of use, and overlap with coral habitat.

Demersal longlines and long‐lined pots ranked

intermediate in terms of potential threat to deep

coral habitats while mid‐water trawls, single

pot sets, and scallop dredges were considered

to pose minimal threat to those habitats. The

effects of other human activities on coral

habitats were discussed with only climate

change and associated effects of ocean

acidification noted as being sources of real

concern. Recent measures to protect coral

habitat via Essential Fish Habitat (EFH) by the

North Pacific Fishery Management Council

were detailed with special reference to Habitat

Areas of Particular Concern (HAPCs) that were

implemented about the time the 2007 Report

was published. Regional priorities to manage

and conserve deep coral communities were

outlined, with specific recommendations on

future research activities that included studies

on growth rates and reproductive ecology,

taxonomy, the effects of ocean acidification and

oil toxicity, the effects of specific fishing gear

types on coral habitat, and a call for

reconnaissance submersible dives in coral

“hotspots” to assess their suitability as HAPCs.

The 2007 Report also called for increased use of

coral bycatch data from fisheries and stock

assessment surveys as a source of mapping

Alaska’s coral resources but noted that this

would require modifications to the way data

were currently being collected and the

implementation of new observer training

programs and publication of new field guides.

This update reviews and highlights new

information since the 2007 report and through

2016.

II. Update on Scientific

Information

II.1. New Research – Overview

Prior to 2012, few major field research efforts

funded directly by NOAA had been initiated

since the publication of the 2007 Report (see

Figure 1). Several small programs, focused

mostly on biological processes, have been

established with limited funding from various

sources. A major research initiative for Alaska

was established for 2012–14 as part of NOAA’s

Deep Sea Coral Research and Technology

Program. Here we report on new knowledge

58
                     

     

               

                 

               

             

           

               

       

               

             

         

             

          

       

         

           

           

             

             

             

               

         

           

         

           

                   

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Figure 1. Location of research projects completed or initiated since 2007.

gained about deep corals (and sponges) in the

past 8 years in Alaska that includes findings for

studies done prior to the 2007 but excluded

from the previous Report (Stone and Shotwell

2007), as well as new resource management

measures, and we provide a new list of

priorities for future research.

A major research program was initiated in 2003

to investigate coral habitat in the central

Aleutian Islands using the manned submersible

Delta and the remotely operated vehicle (ROV)

Jason II. NOAA’s National Marine Fisheries

Service (NOAA/NMFS), the North Pacific

Research Board, and NOAA’s Undersea

Research Program sponsored this research. A

comprehensive report for this research was

completed (Heifetz et al. 2007) with detailed

manuscripts on damage to deep‐sea coral and

sponge habitats (Heifetz et al. 2009), modeling

coral habitats (Woodby et al. 2009), and a

comprehensive manuscript on the ecology of

coral and sponge habitat (Stone 2014).

Greenpeace conducted research cruises to

Pribilof and Zhemchug Canyons along the

59
                     

     

               

             

             

           

               

       

         

             

               

               

               

           

           

           

           

         

           

           

             

               

           

         

       

  

             

               

           

             

           

              

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Bering Sea continental margin in 2007 and 2012

to investigate seafloor habitats (Miller et al.

2012, Miller et al. 2015). NOAA scientists

continue to collaborate with University of

Maine scientists in studying the ecology of red

tree coral (Primnoa pacifica) in shallow‐water

glacial fjord habitats. Additional small‐scale

field studies have been conducted on the

biology of corals and associated species in areas

closed to bottom trawling on the western Gulf

of Alaska slope, rockfish habitat in the eastern

Aleutian Islands and previously trawled areas

in the eastern Gulf of Alaska.

II.1.i – The Alaska Coral and Sponge

Initiative: a NOAA Deep Sea Coral

Research and Technology Program

regional fieldwork initiative in Alaska

From 2012–2015, NOAA’s Deep Sea Coral

Research and Technology Program and Alaska

Fisheries Science Center conducted a field

research initiative in Alaska to better under‐

stand the location, distribution, ecosystem role,

and status of deep‐sea coral and sponge

habitats (Rooper et al. 2017). The initiative was

developed in consultation with the North

Pacific Fishery Management Council and

targeted information that could inform

management.

The Alaska Coral and Sponge Initiative was

comprised of a series of research projects that

began in the summer of 2012:

 One mapping cruise, two ROV cruises, and

one drop‐camera cruise to map areas of

high abundance of Primnoa corals in the

Gulf of Alaska and study the ecological

function of these habitats.

 Two cruises to determine the distribution of

high abundance and diversity areas of

deep‐sea corals and sponges in the Aleutian

Islands through modeling and field

sampling using towed/drift cameras.

 New estimates of the recovery rates and

sustainable impact rate for Primnoa corals in

the Gulf of Alaska through a landscape

ecology approach.

 A field study to determine the productivity

of commercial fishes from coral and non‐

coral habitats in the Gulf of Alaska.  

 A field study to develop and test the

feasibility of using a towed/drift camera

system to estimate the effects of commercial

long‐line and pot fishing on deep‐sea coral

and sponge communities in the Gulf of

Alaska.

 Conducted a genetic study of population

connectivity of red tree corals (Primnoa

pacifica) in the eastern Gulf of Alaska.

 Initiated collection of long‐term data sets of

oxygen and pH from summer bottom trawl

surveys.

 Established a long‐term monitoring station

at a shallow‐water fjord coral site in

Southeast Alaska.  

 Examined existing and newly collected data

and specimens to improve the taxonomy of

deep‐sea corals and sponges and conduct

paleoclimatological, reproductive ecology,

trophic dynamics and marine natural

products studies.  

 Compiled a geologically based substrate

map for the Gulf of Alaska and Aleutian

Islands.  

60
                     

     

             

               

       

               

                 

             

   

         

       

               

         

           

             

         

         

               

             

             

             

               

  

               

                     

               

             

                 

               

             

          

            

     

     

           

        

           

           

         

     

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

In addition to these projects, the Alaska

Fisheries Science Center and the Deep Sea Coral

Research and Technology Program collaborated

on surveys and modeling of corals and sponges

on the eastern Bering Sea slope and canyons at

the request of the North Pacific Fishery

Management Council.

II.2. Taxonomy and Species

Distributions

II.2.i – Corals

a. Coral taxonomy

The 2007 Report listed 141 unique coral taxa

including 11 scleractinians, 14 antipatharians,

15 alcyonaceans (including six stoloniferan

taxa), 63 gorgonians, 10 pennatulaceans, and 28

stylasterids. The list included 52 taxa with

incomplete taxonomy and several taxa that

were only recently collected and awaited

formal description. Since that time, Cairns and

Baco (2007) described five new species of  

Narella from Gulf of Alaska seamounts

previously known only generically and Etnoyer

(2008) described a new species of bamboo coral

(Isidella tentaculum), a major structure forming

coral, previously known only generically from

deepwater areas of the Gulf of Alaska

seamounts and the Aleutian Islands.

Additionally two major revisions were

undertaken – one on the Primnoidae from the

Aleutian Islands and Bering Sea (Cairns 2011)

and another on the Stylasteridae from Alaska

and adjacent waters (Cairns and Lindner 2011)

– both added many new species and eliminated

numerous synonyms.

Advances in coral taxonomy have been made in

Alaska in the past 8 years but there are still a

few species to describe, both in hand and

known from video footage. The updated list

(Stone and Cairns 2017 – Online Annex) now

includes 137 unique coral taxa (Table 1) known

from Alaskan waters but still includes three

dozen species with incomplete taxonomy.

Table 1. Updated summary of species richness and depth range for seven major groups of corals found in

Alaskan waters. Data sources and references are found in the Online Annex 1.

.TAXA Number of Species Depth Range

(m)

Black corals (Order Antipatharia) 12 22–6328

Stony corals (Order Scleractinia) 12 401–4784

Soft corals (Order Alcyonacea, in part)

Suborder Alcyoniina

Suborder Stolonifera

8

6

10–3209

11–591

Gorgonian corals (Order Alcyonacea, in part) 61 6–4784

Sea pens and whips (Order Pennatulacea) 14 3–2947

Lace corals (Family Stylasteridae) 24 10–2124

Total 137 3–6328

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Figure 2. Alaska seascapes: a) Rich sponge and coral habitat on Dickins Seamount in the Gulf of Alaska. b) Red

tree coral (Primnoa pacifica) thicket on Shutter ridge in the Gulf of Alaska. c) Sponge habitat near Fairweather

Ground in the Gulf of Alaska. d) Coral and sponge “garden” in the Aleutian Islands.

Some taxa are still known only generically,

possibly including some synonyms and names

borrowed from other areas and used simply as

placeholders. All major coral groups are in need

of additional taxonomic work but the

Alcyonacea and Pennatulacea are most in need

of major revision. Additionally, the gorgonian

families Plexauridae and Isididae are in need of

taxonomic attention. Specimens are available to

do the work but there is currently a shortage of

taxonomists with expertise in these groups.

b. Coral distribution

The principal source of coral distribution

information in Alaska continues to be catch

data collected during annual research trawl

surveys. Since the survey stations are located at

the same approximate locations each year no

new spatial information is available from that

source when considered at the spatial scale of

the surveys. These trawl surveys have been

extended in recent years to cover new areas in

the arctic and subarctic waters of the northern

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Bering Sea, Chukchi Sea and Beaufort Sea.

Trawl survey data are currently being used to

develop models of coral and sponge

distribution applicable to all Alaska regions

(Rooper et al. 2014, Sigler et al. 2015).  These

trawl surveys have now been complemented by

towed/drift camera surveys conducted in the

Aleutian Islands (Goddard et al. in rev.) and

eastern Bering Sea (Goddard et al. 2016).

Another tremendous source of coral

distribution information is from commercial

fisheries bycatch records maintained by the

Alaska Fisheries Science Center and collected

by observers as part of the Fisheries Monitoring

and Analysis Division. Unfortunately, the

taxonomic resolution presently used by

observers provides little fine‐scale information

on coral distribution. NOAA initiated a pilot

project in 2012 to improve the taxonomic

identification capability of fisheries observers

by providing them with training, a field guide,

and a sampling protocol to test their ability

(Stone et al. 2015).  In general, the success of

sample identification was low but highly

variable. Long‐term implementation of a coral

identification protocol, like the one tested for

this project, would require training resources

beyond those currently available.

There have been few studies of population

connectivity for Alaskan corals.  Baco and

Cairns (2012) concluded that haplotypes for

corals of the genus Narella collected on

seamounts in the Gulf of Alaska and Hawaii

had relatively narrow depth and geographic

ranges.  Analysis of genetic connectivity for

Primnoa pacifica in the eastern Gulf of Alaska is

ongoing and genetic markers have been

developed for this very important genus of

octocorals (Morrison et al. 2015).

Spatial data from major projects completed

prior to the 2007 Report are now available (e.g.

Stone 2014, Stone et al. 2014a,b) and additional

spatial data will also soon be available as part

of another NOAA project to review the

submersible‐collected video archives

throughout Alaska. Coral specimens with

precise location data archived in the Auke Bay

Laboratory (n = ~400) are also currently being

re‐cataloged and will soon be available for

spatial analyses.

Bering Sea: The canyons and slope of the

eastern Bering Sea were a major focus of new

surveys and analysis. Greenpeace conducted a

research cruise to Pribilof and Zhemchug

Canyons along the Bering Sea continental

margin in 2007 (Miller et al. 2012). High

densities of corals (0.43 colonies m‐2) were

reported from the canyons (combined) and

principally included scattered groves of the

pennatulacean Halipteris willemoesi and fields of

the gorgonian Plumarella aleutiana on several

transects. Miller et al. (2012) reported that

Pribilof Canyon supported higher densities of

corals (0.79 colonies m‐2) and sponges (0.53

individuals m‐2) than Zhemchug Canyon (0.07

colonies m‐2 and 0.01 individuals m‐2,

respectively). Most notably, the study

documented several northern range extensions

and new records for the region: 1) the

scleractinian Caryophyllia alaskensis (a northern

range extension); 2) the antipatharian Lillipathes

wingi (northernmost record in the Pacific

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Figure 3. Deep‐sea corals of Alaska: a) Large bubblegum coral (Paragorgia sp.) from Pratt Seamount in the Gulf

of Alaska. b) The gorgonian Fanellia fraseri in a bed of scallops, many encrusted with the sponge Myxilla

parasitica in the Aleutian Islands. c) Gorgonian coral (Isidella tentaculum) with two galatheid crabs from

Welker Seamount in the Gulf of Alaska. d) Black coral (Parantipathes sp.) at a depth of 1003 m in the Aleutian

Islands. e) Stylasterid coral (Stylaster campylecus) at a depth of 681 m in the Aleutian Islands. f) Sharpchin

rockfish in a large Primnoa pacifica in the Gulf of Alaska.

64
                     

     

     

           

           

                 

   

         

       

               

           

             

                   

         

             

               

           

               

         

               

           

             

                   

             

             

               

                 

               

             

             

               

           

           

             

               

           

                 

             

             

               

         

             

               

               

             

               

                     

             

               

                   

               

             

           

                 

             

           

             

             

             

               

               

   

             

               

              

           

             

           

         

               

        

           

             

                  

     

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Ocean); 3) the alcyonacean Heteropolypus sp.

(=Anthomastus sp.; likely a northern range

extension for the genus); 4) a stoloniferan

Clavularia sp. (a new record for the region); 5)

six gorgonians (Plumarella aleutiana, Primnoa

pacifica, P. wingi, Keratoisis sp., Swiftia pacifica,

and Paragorgia arborea) all representing new

range extensions or new records for the region;

and 6) two pennatulaceans (Protoptilum sp. and

cf. Pennatula sp.), each representing a range

extension.

Based on the findings of the Miller et al. (2012)

study, the North Pacific Fishery Management

Council requested analyses (Sigler et al. 2015,

see below) and a model validation survey to

determine if coral concentrations in two

canyons in the eastern Bering Sea, Pribilof and

Zhemchug, warranted further protection from

fishing activity. The Council asked the AFSC to

collect information on coral presence inside and

outside the canyons, information on the height

and density of corals, and the role of corals as

fish habitat. They also asked scientists to

document the presence and degree of fishing

gear effects. The field survey was conducted in

2014 and the results were reported in Rooper et

al. (2016), Goddard et al. (2016), and MacLean

et al. (2017). A towed/drift stereo camera

system was deployed at 250 randomly selected

sites along the eastern Bering Sea shelf and

outer slope. The results validated previous

modeling and analysis work, confirming that

coral habitat occurs both inside these two

canyons and along the Bering Sea slope. In

general, coral densities throughout our survey

area were low where they occurred. This is not

surprising as the eastern Bering Sea seafloor

contains little (~2.8% of the total observed

seafloor) of the rocky habitat that most corals

require for attachment. The highest

concentration of coral habitat was found in

Pribilof Canyon and to the northwest along the

slope. Sea whip densities were highest in sandy

portions of the slope through‐out our survey

area. Most gorgonian corals observed were < 30

cm tall, while sea whips were as tall as 1.5 m.

Direct evidence of fishing (mostly trawl tracks,

but also derelict gear) was observed at 32

(12.8%) of the sample sites. In total, 2.9% of the

corals and 0.3% of the sponges observed were

damaged. About 9% of individual sea whips

observed were classified as either damaged,

dead or lying horizontal on the seafloor. It was

difficult to determine if the damage was

human‐induced (e.g., fishing or other activity)

or natural (e.g., sea star predation, ocean

currents). There were very few places where

both clear evidence of fishing activity (e.g.,

trawl tracks or fishing gear) and damaged coral,

sponges or sea whips occurred (3.2% of the

camera transects).

Aleutian Islands: The 2007 report identified the

Aleutian Islands as the region with the highest

abundance and diversity of corals in Alaska.

Stone (2014) presented analyses of deep‐sea

coral and sponge habitats from video surveys

conducted from 2002–2004 in the central

Aleutian Islands, greatly expanding the

geographic and depth (to 2947 m) range of

earlier observations. Corals were widely

distributed throughout the study area, with the

highest density observed at depths between 400

– 700 m. An additional 18 coral and sponge

gardens were identified.

65
                     

     

           

           

               

             

             

           

                 

       

           

               

             

                 

             

             

               

           

           

           

           

             

       

       

               

             

             

             

               

     

             

               

               

           

           

             

             

           

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

In 2012 and 2014, camera surveys were

conducted at 216 randomly‐selected sites

throughout the Aleutian Islands and Bowers

Ridge, using protocols similar to those used on

the eastern Bering Sea surveys.  These surveys

indicated that high‐density coral and sponge

communities were much more extensive than

previously surveyed (Goddard et al., 2017).  

Survey data were used to ground‐truth the

coral and sponge distribution models

developed for the Aleutian Islands (Rooper et

al. 2014; see below).    

Gulf of Alaska:  In August 2010, there was an 8‐

day cruise aboard the chartered fishing vessel

Sea Storm to investigate three slope areas closed

to bottom trawling in 2007 in the western Gulf

of Alaska. These slope areas (Shumagin, Sanak,

and Unalaska) were identified by fishers to

potentially contain coral and sponges in

virtually pristine states, as fishing has not

typically occurred in these areas. The objective

of the cruise was to use video cameras to

determine whether corals and sponges

occurred in each of the three areas and to collect

data that would allow predictive modeling of

their distribution there. Upright sponges (vase

or arborescent morphology) occurred at all

three closed areas, although they were in

relatively low abundances at the Unalaska and

Shumagin areas.  Deepwater corals were not

found at the Shumagin area and were observed

only at one site in Unalaska. Corals occurred at

14 of 30 sites in the Sanak area. Sponges were

distributed across all depths where video was

recorded, while corals were distributed only at

the shallowest locations (< 150 m) in the closed

areas. The corals observed were most

commonly upright gorgonians with a fan‐type

morphology. Corals were present only where

boulders were present and did not occur in

areas with only sand and gravel substrates.

Sponges occurred where both boulders and soft

mud substrates were present, although they

were also more likely to occur at locations with

cobble or boulders present.

Areas supporting dense red tree coral (Primnoa

pacifica) thickets in the eastern Gulf of Alaska,

including the five small areas designated as

HAPCs by the NPFMC in 2006, have been focal

sites for studying some of Alaska’s most

important coral habitats. Extensive in situ work

conducted there in 2005 (Stone et al. 2014a)

revealed that seafloor habitats are dominated

by red tree corals, hydrocorals (Stylasteridae), a

few other octocorals (Calcigorgia spp., Paragorgia

sp., and Halipteris willemoesi) in lower numbers,

demosponges and in some areas dense sponge

grounds of hexactinellids (mainly Aphrocallistes

vastus, Heterchone calyx, and Farrea occa). The

presence of red tree corals is highly associated

with bedrock, high seabed roughness and slope

(Masuda and Stone 2015) and the habitats

formed by these corals and sponges provide

essential habitat for some species of fish (Stone

et al. 2014a).

Alaskan Arctic: New surveys conducted in the

Chukchi Sea in 2009 and 2010 in association

with proposed oil and gas leases (BOEM, 2012)

revealed a patchy distribution of habitats

structured by sedentary invertebrates. The soft

coral (Gersemia sp.) was the primary coral

species identified, with more diverse and dense

coral and sea anemone communities at

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

nearshore sites and within Barrow Canyon.

These findings were confirmed during a cruise

in July 2012 led by Greenpeace.

II.2.ii – Sponges

a. Sponge taxonomy

NOAA’s Deep Sea Coral Research and

Technology Program provided funding for the

publication of “A Guide to the Deepwater

Sponges of the Aleutian Islands Archipelago”

(Stone et al. 2011). The main purpose of the

guide was to promote an awareness and

appreciation of the importance of the sponge

fauna in the North Pacific Ocean, particularly in

the Aleutian Islands where the diversity and

abundance of sponges appear to be

extraordinary and where bycatch in existing  

fisheries continues to be a major concern for

resource managers. The guide documented 196

sponge taxa from deep‐water (> 80 m) in

Alaskan waters (Table 2), but noted that the

inventory is largely incomplete and that there

are many species yet to be discovered in the

region. Reiswig and Stone (2013) further

described deep‐water hexactinellid sponges

from the central Aleutian Islands, including a

new genus and eight new species in five

families. A special collections project was

initiated in 2012 to collect select sponge taxa in

the Aleutian Islands aboard the biennial

research trawl survey conducted there by

NOAA. Nearly a dozen new species of sponges

have already been discovered as part of this

project (Lehnert et al. 2013; Lehnert and Stone

2014a, 2014b; Lehnert and Stone 2015).

Table 2. Summary of species richness and depth range for classes of sponges found in Alaskan waters, based

on Stone et al. (2011) and Reiswig and Stone (2013).

CLASS

Number

of

Species

Depth Range

(m)

Calcarea 12 5–250

Hexactinellida 52 20–2800

Demospongiae 130 0–2800

Homoscleromorpha 2 95–383

Total 196 0 – >2800

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Figure 4. Deep‐sea sponges of Alaska: a) Hexactinellid sponge Farrea occa occa at a depth of 165 m in the

eastern Gulf of Alaska. b) Glass sponge Heterochone calyx calyx at a depth of 181 m in the eastern Gulf of

Alaska. c) Demosponge Guitarra abbotti at a depth of 146 m in the central Aleutian Islands. d) Demosponge

Mycale (Mycale) loveni at a depth of 96 m in the central Aleutian Islands. e) Demosponge M. loveni with a

gravid sharpchin rockfish (Sebastes zacentrus) at a depth of 170 m in the eastern Gulf of Alaska. f) Glass sponge

Aphrocallistes vastus with a juvenile golden king crab (Lithodes aequispina) in pre‐molt condition at a depth

of 190 m in the central Aleutian Islands.

68
                     

     

b. Sponge distribution

             

               

              

             

                 

             

             

               

           

            

                 

              

             

             

                 

   

       

                 

                 

                 

           

         

           

                 

             

                 

             

         

           

           

       

     

  

             

           

           

               

             

                 

             

             

           

             

           

           

         

           

          

           

             

         

         

                  

               

             

          

               

                 

           

             

               

             

             

               

               

                    

                 

             

             

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Sponges are a major component of biogenic

habitat or “living substrate” in the Gulf of

Alaska, Aleutian Islands and the Bering Sea.

Sponges are the most common taxon forming

living substrate in the Gulf of Alaska and the

Aleutian Islands (Malecha et al. 2005). Sponges

reach high densities in the Aleutian Islands

(Stone et al. 2011), and demosponges are a

major component of the region’s coral and

sponge “gardens” (Stone 2014). Sponges are

also common in the Bering Sea, with some areas

of very high density in Bristol Bay. Sponges

have also been reported from the Alaskan

Arctic (de Laubenfels 1953, BOEM 2012), but

appear to have occurred in low numbers at the

sites sampled.

Bycatch in fisheries and fisheries‐independent

surveys is a major source of information on the

location of the sponge fauna, and a source of

specimens for study. The guide by Stone et al.

(2011) served the additional purpose of

providing fisheries observers and scientists

with the information necessary to adequately

identify sponge fauna so that the data can be

included in existing databases. These data can

be used to map areas of high abundance and

the locations of indicator species of vulnerable

marine ecosystems. The guide was also

designed for use by scientists making

observations of the fauna in situ, with

submersibles including remotely operated

vehicles (ROVs) and autonomous underwater

vehicles (AUVs).

A research cruise (March 2010) in Portland

Canal along the border between Southeast

Alaska and British Columbia, confirmed the

presence of small sponge reefs detected a few

years previously with sonar by the Geological

Survey of Canada (Stone et al. 2014b). The reefs

were similar in species composition to, but

much smaller in size than, the massive

bioherms reported farther south in British

Columbia (Conway et al. 2005). Small patches

of hexactinellid sponge reef, believed to be

biohermal, were also observed in northern

Southeast Alaska near Juneau, possibly

indicating that these structures are more

common than originally hypothesized.

II.2.iii – Modeling distribution and

abundance of corals and sponges

Predictive habitat modeling for deep‐sea corals

and sponges provides an important tool for

understanding the distribution of these

organisms across Alaska’s huge geographic

regions. Since the 2007 report, there has been a

significant expansion in the use and utility of

predictive habitat models in Alaska (Guinotte et

al., this report). These include presence‐only

models (Maxent) of coral taxa (grouped at the

Order or suborder level) for the Gulf of Alaska,

Aleutian Islands and eastern Bering Sea

(Guinotte and Davies 2013); models for coral

gardens in the Aleutian Islands (Woodby et al.

2009; Guinotte and Davies 2013); and models

for coral and sponge presence, abundance and

diversity in the Aleutian Islands (Rooper et al.

2014) and eastern Bering Sea slope and canyons

(Sigler et al. 2015, Rooper et al. 2016). Rooper et

al. (2016) presented one of the first examples of

rigorous validation of coral and sponge habitat

models, using camera surveys to assess the

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

accuracy of predictions from a model based on

trawl surveys along the eastern Bering Sea

slope.   

The results of these modeling studies supported

observations on the relative richness of coral

and sponge habitats in the Aleutian Islands

compared to most other regions, and predicted

that these habitats were expected to be much

more widespread in the Archipelago.   The

North Pacific Fishery Management Council

utilized model results in its deliberations on

potential protections for Bering Sea canyons.

II.3. Species Associations with

Deep-Sea Corals and Sponges

In Alaska, commercial fisheries species are

associated with deep corals and sponges at both

fine‐scales (less than 10 m, e.g. Stone 2006) and

at the scale of existing fisheries (e.g. Heifetz

2002). Research conducted since or not reported

in the 2007 Report continue to support earlier

observations that fish and crabs, particularly

juveniles, are associated with coral habitat

throughout Alaska and are presumably using it

as refuge and as focal sites of high prey

abundance (Rooper and Boldt 2005; Rooper et

al. 2007; Rooper and Martin, 2012; Miller et al.

2012; Stone 2014; Stone et al. 2014a). A study in

the eastern Aleutian Islands found juvenile

Pacific ocean perch used these habitats as

nursery areas and were closely associated with

boulders and rocky areas that were extensively

covered with coral and sponge (Rooper et al.

2007). All of these studies have measured

association at varying scales and have utilized

different methods to test association but are

noteworthy since these observations are not

well quantified in many other regions and

habitats where deep‐sea corals occur. Not all

demersal species are associated with coral

habitat in Alaska. For example, Rooper and

Martin (2009) found that shortspine thornyhead

(Sebastolobus alascanus) catches were not

associated with catches of corals and sponges in

the Gulf of Alaska bottom trawl survey. Stone

(2014) also found that some deep‐water species

in the Aleutian Islands are no more likely to

occur with emergent epifauna than they are

without emergent epifauna.

II.4. Ecological Studies (Growth,

Reproductive Ecology, Microbial

Ecology, and Marine Natural

Products)

New studies on growth rates of Alaskan corals

corroborated earlier observations from Alaska

and elsewhere that they are slow‐growing and

consequently long‐lived. Andrews et al. (2009)

used 210Pb techniques to determine the growth

rate for two species of bamboo corals in the

Gulf of Alaska (~1.4 cm y‐1 axial growth rates

for Isidella tentaculum and ~1.0 cm y‐1 for

Keratoisis sp.). Stone et al. (2017) analyzed the

growth of tagged colonies of the shallow‐water

holaxonian Calcigorgia spiculifera in Southeast

Alaska over a period of five years. The growth

rate of 0.6 cm y‐1 was the slowest rate reported

for any gorgonian in Alaskan waters. Andrews

and Stone (unpublished data) used growth ring

counts to estimate a growth rate between 1.2–

1.8 cm y‐1 for the primnoid octocoral Fanellia

compressa and Stone (unpublished data) used a

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

deglaciation record to estimate a growth rate

for Primnoa pacifica that validated the rate

previously established for the species (Andrews

et al. 2002).  

A study was undertaken on the reproductive

ecology of stylasterid corals in the Aleutian

Islands (Brooke and Stone 2007) that indicated

that they are gonochoristic brooders with

limited potential to provide sources of recruits

to disjunct disturbed habitats. Studies on the

reproductive ecology and ultrastructure of P.

pacifica are being undertaken in shallow‐water

populations in the Tracy Arm fjords of

Southeast Alaska (Waller et al. 2014) and

should provide insights into the recovery

dynamics of this important species.     

The microbial ecology of corals was studied in

both the Aleutian Islands (Gray et al. 2011) and

on Gulf of Alaska seamounts (Penn et al. 2006).

Both studies indicate that corals harbor unique

and diverse bacterial communities that may

provide the corals with a wide array of benefits

including the facilitation of processes for a

portion of their nutrition. The Alaska Fisheries

Science Center and University of Mississippi

have been routinely collecting sponges from

survey trawls in the Aleutian Islands since 2004

and investigating their use as marine natural

products, specifically as biomedicines. Dozens

of sponge taxa have been investigated to date

and a few have shown significant promise as

antiviral, antimalarial, and antimicrobial agents

(Na et al. 2010). Additional collections of

demosponges from the Aleutian Islands are

being analyzed and one species has already

yielded extracts with evidence of bioactivity

against opportunistic infectious diseases,

malaria, and Hepatitis C (Abbas et al. 2011).

II.5. Research Priorities and

Planned or Anticipated Research

Activities

In preparation for the upcoming fieldwork

under NOAA’s Deep Sea Coral Research and

Technology Program, a workshop was held in

Anchorage in September 2010 to identify

research priorities for the region, including

critical information needs and research

activities to address these needs (NOAA 2010).

Key priorities included the following:

 Expanding our knowledge of the

distribution of deep‐sea corals and sponges

and their habitats by analyzing existing

information, new seafloor mapping and

visual surveys, and by developing

predictive habitat models. A geographic

priority was placed on the Gulf of Alaska

and Aleutian Islands.

 Determining the population characteristics

of deep‐sea corals and sponges, including

growth rates, life history and reproductive

traits, trophic dynamics, environmental

tolerances, and population connectivity.

 Determining the key functions that deep‐

sea corals and sponges provide to managed

species at different life stages.

These priorities were largely derived from

ongoing research needs and objectives

identified by the Program, the North Pacific

Fishery Management Council, and the Essential

Fish Habitat‐Environmental Impact Statement

process. They formed the basis of the 2012–2014

Alaska Coral and Sponge Initiative (Rooper et

al. 2017).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

III. Update on Management

of Deep-Sea Corals and

Sponges

III.1. New Information on Impacts

and Stressors

III.1.i – Fishing

The 2007 Report indicated that the major

stressor on deep‐sea coral and sponge

communities in Alaska was the impact of

commercial bottom‐fishing activities and new

information continues to support this premise

(Heifetz et al. 2009, Stone 2014, Rooper et al.,

this volume). Accordingly, studies on the

effects of fishing on seafloor habitat continue to

be a priority in Alaska with particular emphasis

on the recovery rates of disturbed habitats.

Studies were implemented to examine the

recovery rate of sponges 13 years post‐

(experimental) trawling in low density sponge

habitat in the eastern Gulf of Alaska (Malecha

and Heifetz, unpublished data) and recovery

from simulated trawl disturbance on the

pennatulacean Halipteris willemoesi (Malecha

and Stone 2009) and gorgonian Calcigorgia

spiculifera (Malecha et al., unpublished data). In

2005, scientists used the submersible Delta to

document ambient levels of disturbance to

corals and sponges in eastern Gulf of Alaska

Primnoa thickets set aside as HAPCs (Stone et

al. 2014a). The HAPCs have been in an area

closed to bottom trawling since 1998 so

observations of disturbance represent those

principally from longline fishing. Rooper et al.

(2011) modeled the impacts of bottom trawling

in the Aleutian Islands and subsequent

recovery rates of corals and sponges and

predicted recovery rates were slow, especially

for corals, where the coral biomass would

require decades to recover.

As with corals, the primary stressor on

deepwater sponge communities in Alaska is

likely physical disturbance by commercial

bottom‐contact fishing gear. The first National

Bycatch Report (NMFS 2011) identified an

average annual bycatch in 2003–2005 of over

525,000 lbs. (238,000 kg) of sponges. Bottom

trawl fisheries in the eastern Bering Sea and

Aleutian Islands fisheries had the highest

sponge bycatch. Despite extensive bottom‐

trawl closures in the Aleutian Islands since

2006, updates to the National Bycatch Report

(NMFS 2013, 2016) showed continued high

levels of sponge bycatch in certain fisheries

from 2010–2013.

III.1.ii – Other stressors

The effects of other human activities on coral

habitats–deployment of pipelines and cables,

point‐source pollution discharges, fish

processing waste disposal, mineral mining, and

invasive species–continue to be of minimal or

nonexistent concern, although interest in

offshore petroleum drilling in the arctic regions

of Alaska may present concerns regarding the

presence of soft corals there. The Bureau of

Ocean Energy Management (BOEM) approved

a Shell Oil Chukchi Sea Exploration Plan;

however, Shell Oil subsequently announced in

September 2015 that it would cease exploratory

offshore drilling in the Arctic. Climate change

and the associated effects of ocean acidification

on coral and sponge habitats continue to be

sources of real concern, as discussed below.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

III.1.iii – Climate change and ocean

acidification

Decreases in oceanic pH and resulting

decreases in calcium carbonate saturation state

could have profound effects on corals

dependent on the extraction of calcium

carbonate from seawater for skeletal building.

Corals will be affected differently depending on

their skeletal composition (aragonite vs. calcite

vs. magnesium calcite), geographical location,

and depth relative to the already particularly

shallow calcium carbonate saturation horizons

in the North Pacific Ocean.

The skeletal composition is unknown for most

species of deep‐sea corals worldwide and is

known for only a handful of the 137 taxa

presently documented from Alaskan waters. To

that end, a study was initiated in 2010 by

sorting through the extensive archives at the

Auke Bay Laboratories and Smithsonian

Institution; 130 specimens comprising 61 taxa

from all major groups of corals (scleractinians,

gorgonians, true soft corals, stoloniferans,

pennatulaceans, and stylasterid corals) were

selected for laboratory analyses. Multiple

specimens were selected for taxa of particular

ecological importance (i.e. those that form large

single‐species assemblages), and specimens of

the same species from multiple depth and

geographic zones.

Laboratory analyses were performed at the

Department of Geology and Geophysics at the

Woods Hole Oceanographic Institution and the

Center for Material Sciences and Engineering at

the Massachusetts Institute of Technology. X‐

ray diffraction and full‐pattern Rietveld data

refinement were used to precisely determine

the skeletal composition of Alaskan corals.

Corals composed of high magnesium‐calcite are

the most soluble. Consequently, those corals,

particularly those residing at depths deeper

than the saturation horizon, are most at risk to

decreases in oceanic pH unless they have

adapted physiological processes to counter the

effects.

The mineralogy data will be used in

conjunction with species distribution data

(depth and geographical) and the present and

projected aragonite and calcite saturation

horizons in Alaska to predict the effects of

ocean acidification on coral resources of the

North Pacific Ocean. At the completion of this

project a comprehensive risk assessment will be

conducted for all corals in Alaskan waters but

physiological data are still needed for Alaskan

coral taxa, probably in a laboratory setting, to

determine if taxa respond as predicted by

mineralogy and marine chemistry data.

Principal Investigators of this research are

Robert Stone (NOAA, AFSC), John Guinotte

(Marine Conservation Institute), Angela

Helbling and Anne Cohen (Woods Hole

Oceanographic Institution), and Stephen Cairns

(Smithsonian Institution).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

III.2. New or Planned

Management Actions

III.2.i – Fishing

A suite of closures were approved by the North

Pacific Fishery Management Council in 2005,

including the Aleutian Islands Habitat

Conservation Area (AIHCA), and implemented

on 28 July 2006. Since that time, five additional

management activities involving spatial

closures that potentially harbor deep coral and

sponge habitat have been approved and

implemented by the Council and NOAA.

Effective 20 March 2008, Amendment 88 to the

Fishery Management Plan for Groundfish of the

Bering Sea and Aleutian Islands Management

Area revised the boundaries of the AIHCA to

allow bottom trawling in a historically fished

area (north of Agattu Island) and to prohibit

bottom trawling in an area known to harbor

coral and sponge habitat (west of Buldir

Island). This action was intended to ensure that

the intentions of the AIHCA to protect areas of

coral and sponge habitat from the potential

effects of bottom trawling were realized and to

allow non‐pelagic trawling to continue in areas

Figure 5. Commercial fishing closures in Alaska (data provided by John Olson, NMFS‐Alaska Regional

Office, Anchorage, Alaska).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

historically fished. The opening of the area

north of Agattu Island potentially puts coral

and sponge habitat at risk since it has not yet

been surveyed for coral and sponge habitat.  

The Northern Bering Sea Research Area was

implemented in 2008 and prohibited bottom

trawling in a broad region there. The intention

of the closure was to provide an opportunity to

study the potential effects of bottom trawling

on seafloor habitat before authorizing that

activity there. The benthic fauna of the region is

neither well studied nor known to harbor much

coral or sponge habitat so the conservation

relevance to these habitats is unknown.  

In 2009, a new Arctic Fishery Management Plan

for the Fish Resources of the Arctic

Management Area (Chukchi and Beaufort Seas)

was implemented, essentially closing all federal

waters in the region to commercial fishing

activities. The Council was concerned that

changing ecological conditions could lead to the

development of inadequately regulated

commercial fisheries in the region that could

have adverse effects including those on fish

habitat. The benthic fauna of the region is not

well studied and is not known to harbor much

coral or sponge habitat, so the conservation

relevance to these habitats is unknown.

In 2011, a trawl sweep modification

requirement (Amendment 94) was

implemented for vessels participating in the

Bering Sea flatfish fishery that occurs

exclusively on the broad and homogeneous

continental shelf. The modification requires

discs or bobbins providing 1–3ʺ lift on the trawl

sweeps to limit adverse effects on low‐relief

seafloor biota. This gear modification has been

demonstrated to reduce impacts on patches of

soft corals and sponges located in those habitats

(Rose et al. 2013).

Effective 1 January 2011, trawling and

longlining for two major forage species (Atka

mackerel and Pacific cod) of endangered Steller

sea lions were prohibited in the western

Aleutian Islands. Concurrently, additional

restrictions in 3‐nautical mile (5.56 km) buffer

areas around established sea lion rookeries

went into effect in the central and eastern

Aleutian Islands. The closures were not directly

intended to protect seafloor habitat but will

clearly provide sanctuary to coral and sponge

habitat from bottom trawling in those areas.

The Council also examined whether habitat

protections were warranted to conserve deep‐

sea corals in the Eastern Bering Sea canyons

and slopes. Based on analyses by NOAA (e.g.,

Rooper et al. 2016), the Council concluded in

2016 that deep‐sea corals in these areas are not

at risk from fishing, and additional habitat

protections were not necessary (MacLean et al.

2017).

NOAA is examining the effectiveness of the

AIHCA to protect sensitive seafloor habitat

from the damaging effects of fishing activities.

The approach utilizes a geographic information

system (GIS) to examine patterns of coral and

sponge bycatch abundance and fishing

intensity before and after the closures with

particular reference to areas that have remained

open to fishing.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

III.2.ii – Other management actions

In 2016, the Department of Interior announced

final regulations to ensure that future

exploratory drilling activities on the Outer

Continental Shelf within U.S. waters of the

Beaufort and Chukchi Seas will be conducted

under the highest safety and environmental

standards and subject to strong and proven

operational requirements.

IV. Conclusions

Research on corals conducted in Alaska since

2007, and particularly since 2012 with extensive

new surveys under NOAA’s Deep Sea Coral

Research and Technology Program, continue to

indicate that the region is home to diverse

ecosystems supported by extraordinary coral

and sponge resources. The resources appear to

provide important habitat to fisheries of

national and international importance. There

are still many important gaps in our knowledge

of these ecosystems and analysis continues.

V. Acknowledgements

We thank Jon Heifetz, Wayne Palsson, and John

Olson for providing helpful reviews of this

manuscript. John Olson provided up‐to‐date

background information on management

activities and constructed Figure 5.

Online Annex – Comprehensive list of deep‐

sea corals in the U.S. Alaska region linked here:

https://deepseacoraldata.noaa.gov/library/2017‐

state‐of‐deep‐sea‐corals‐report

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Heifetz J, Woodby D, Reynolds JR, Stone RP (2007)

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structure of the Patton Seamount (Gulf of

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Kelly M, Sim‐Smith C, Stone R, Samaii T, Reiswig H,

Austin W (2016) New taxa and arrangements

within the family Latrunculiidae

(Demospongiae, Poecilosclerida). Zootaxa

2121:1–48

Lehnert H, Hocevar J, Stone RP (2008) A new species

of Aaptos (Porifera, Hadromerida, Suberitidae)

from Pribilof Canyon, Bering Sea, Alaska.

Zootaxa 1939:65–68

Lehnert H, Stone RP (2010) Craniella sputnika sp. nov.

(Porifera: Spirophorida: Tetillidae) from the

Aleutian Islands, Alaska, with suggested

nomenclatural changes for the genera Tetilla and

Craniella. J Mar Biol Assoc U K 91:321‐328

Lehnert H, Stone RP (2014a) Two new species of

sponges from the Aleutian Islands (Porifera,

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Lehnert H, Stone RP (2014b) Aleutian Ancorinidae

(Porifera, Astrophoridae): description of three

new species from the genera Stelletta and

Ancorina. Zootaxa 3826:341–355

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(Porifera, Demospongiae) from the Aleutian

Islands and Gulf of Alaska. Zootaxa 4033:451–

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sp. nov. (Porifera, Demospongiae, Astrophorida)

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Lehnert H, Stone R, and Heimler W (2005a) A new

species of Polymastia (Porifera, Hadromerida,

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Alaska, USA. Facies 51:53–56

Lehnert H, Stone R, Heimler W (2005b) Two new

species of Plakina Schulze, 1880 (Porifera,

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Lehnert H, Stone RP, Heimler W (2006a) New

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Lehnert H, Stone R, Heimler W (2006c) Erylus

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MacLean SA, Rooper CN, Sigler MF (2017) Corals,

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canyons as coral and sponge habitat on the

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loci in the red tree corals Primnoa resedaeformis

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sponge of the genus Latrunculia. J Nat Prod

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Reynolds JR, Rooney SC, Heifetz J, Green HG,

Norcross BL, Shotwell SK (2012) Habitats and

demersal fish communities in the vicinity of

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Geomorphic Features and Benthic Habitat.

Elsevier, London, UK. p 539–553

Rooper CN, Bolt JL (2005) Distribution of Pacific

ocean perch Sebastes alutus in the Aleutian

Islands in relation to benthic habitat. Alsk Fish

Res Bull 11:102–112

Rooper CN, Boldt JL, Zimmermann M (2007) An

assessment of juvenile Pacific ocean perch

(Sebastes alutus) habitat use in a deepwater

nursery. Estuarine Coastal Shelf Sci 75:371–380

Rooper CN, Martin MH (2009) Predicting presence

and abundance of demersal fishes: a model

application to shortspine thornyhead

Sebastolobus alascanus. Mar Ecol Prog Ser

379:253–266

Rooper CN, Martin MH (2012) Comparison of

habitat‐based indices of abundance with fishery‐

independent biomass estimates from bottom

trawl surveys. Fish Bull 110:21–35

Rooper CN, Sigler MF, Goddard P, Malecha P,

Towler R, Williams K, Wilborn R, Zimmermann

M (2016) Validation and improvement of species

distribution models for structure‐forming

invertebrates in the eastern Bering Sea with an

independent survey. Mar Ecol Prog Ser 551:117–

Rooper CN, Stone RP, Etnoyer P, Conrath C,

Reynolds J, Greene HG, Williams B, Salgado E,

Morrison C, Waller R, Demopoulos A (2017)

Deep‐Sea Coral Research and Technology

Program: Alaska Deep‐Sea Coral and Sponge

Initiative Final Report. NOAA Tech. Memo.

NMFS‐OHC‐2, 65 p.

Rooper CN, Etnoyer PJ, Stierhoff KL, Olson JV (this

volume) Effects of Fishing Gear on Deep‐Sea

Corals And Sponges in U.S. Waters. In:

Hourigan TF, Etnoyer PJ, Cairns SD (eds.). State

of Deep‐Sea Coral and Sponge Ecosystems of

the United States. NOAA Technical

Memorandum NMFS‐OHC‐4, Silver Spring, MD

Rooper CN, Wilkins ME, Rose CS, Coon C (2011)

Modeling the impacts of bottom trawling and

the subsequent recovery rates of sponges and

corals in the Aleutian Islands, Alaska. Cont Shelf

Res 31:1827–1834

Rooper CN, Zimmermann M, Prescott MM,

Hermann AJ (2014) Predictive models of coral

and sponge distribution, abundance and

diversity in bottom trawl surveys of the

Aleutian Islands, Alaska. Mar Ecol Prog Ser

503:157–176

Rose C, Munk E, Hammond C, Stoner AW (2010)

Cooperative research to reduce the effects of

Bering Sea flatfish trawling on seafloor habitat

and crabs. AFSC Quarterly Report Feature (Jan‐

Feb‐Mar 2010), 6 p

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Rose CS, Hammond CF, Stoner AW, Munk JE,

Gauvin JR (2013) Quantification and reduction

of unobserved mortality rates for snow,

southern Tanner, and red king crabs

(Chionoecetes opilio, C. bairdi, and Paralithodes

camtschaticus) after encounters with trawls on

the seafloor. Fish Bull 111:42–53

Sigler MF, Rooper CN, Hoff GR, Stone RP,

McConnaughey RA, Wilderbuer TK (2015)

Faunal features of submarine canyons on the

eastern Bering Sea slope. Mar Ecol Prog Ser

526:21–40

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 (2014) The ecology of deep‐sea coral and

sponge habitats of the central Aleutian Islands

of Alaska. U.S. Dep Commer, NOAA

Professional Paper NMFS 16:1–52

Stone, RP, Cairns SD (2017) Deep‐sea coral taxa in

the Alaska Region: depth and geographical

distribution. Online resource:

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Stone RP, Conway KW, Csepp DJ, Barrie JV (2014b)

The Boundary Reefs: glass sponge (Porifera:

Hexactinellidae) reefs on the international

border between Canada and the United States.

U.S. Dep Commer, NOAA Tech Memo

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Stone, RP, Guinotte JM, Helbling A, Cohen AL,

Cairns SD, Cross J (In review) Potential effects of

ocean acidification on Alaskan corals based on

calcium carbonate mineralogy and other factors

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the deep‐water sponges of the Aleutian Island

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year, in situ growth study on shallow‐water

populations of the gorgonian octocoral

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Ecosystems of the United States. U.S. Dep

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of a pilot study to collect coral bycatch data from

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Williams B, Risk M, Stone RP, Sinclair DJ, Ghaleb B

(2007) Oceanographic changes in the North

Pacific Ocean over the past century recorded in

deep‐water gorgonian corals. Mar Ecol Prog Ser

335:85–94

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modeling of coral distribution in the Central

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Online Annex 1. Deep-sea Coral Taxa in the U.S. Alaska Region: Depth and 

Geographic Distribution 

82
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Appendix - Distribution maps for deep-sea corals and sponges in 

Map 1. Locations of black corals (Order Antipatharia) recorded in the National Deep‐Sea Coral and Sponge Database (as o

83
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Map 2. Locations of stony corals (Order Scleractinia) recorded in the National Deep‐Sea Coral and Sponge Database (as of 

84
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Map 3. Locations of true soft corals (Order Alcyonacea) recorded in the National Deep‐Sea Coral and Sponge Database (as 

85
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Map 4. Locations of gorgonian corals (Order Alcyonacea, in part; formerly Gorgonacea) recorded in the National Deep‐SeDatabase (as of November 2017).

86
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Map 5. Locations of sea pens (Order Pennatulacea) recorded in the National Deep‐Sea Coral and Sponge Database (as of N

87
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Map 6. Locations of lace corals (Class Hydrozoa, Order Anthoathecata, Family Stylasteridae) recorded in the National DeeSponge Database (as of November 2017).

88
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Map 7. Location of demosponges (Phylum Porifera) recorded in the National Deep‐Sea Coral and Sponge Database (as of N

89
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Map 8. Location of glass sponges (Class Hexactinellida) recorded in the National Deep‐Sea Coral and Sponge Database (as 

90
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE ALASKA REGION

  

Map 9. Location of calcareous (Class Calcarea) and unidentified sponges (primarily from trawl surveys) recorded in the N

and Sponge Database (as of November 2017).

91
       

         

     

     

   

Large red tree coral

caught as bycatch in a

bottom trawl survey

off Alaska.
EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •

93

EFFECTS OF 

FISHING GEAR ON 

DEEP-SEA CORALS 

AND SPONGES IN 

U.S. WATERS

I. Introduction

Since the 1996 amendment of the Magnuson-Stevens 

Fishery Conservation and Management Act (MSA), 

there have been extensive efforts to identify and protect 

essential fish habitat (EFH). The 2007 MSA reauthorization 

further provided discretionary authority to protect areas 

containing deep-sea corals, recognizing these corals in 

their own right. Beginning around 2005-2007, management 

authorities in some regions of the U.S. acted to protect 

deep-sea corals and sponges. These species are slowgrowing (Leys and Lauzon 1998, Stone and Wing 2001, 

Andrews et al. 2002, Prouty et al., this volume), comprise 

habitat for other species, and may take a long time to 

recover from damage (Gutt and Piepenburg 2003, Rooper 

et al. 2011). They are often damaged by mobile bottomcontact fishing gears, such as bottom-trawls (Chuenpagdee 

et al. 2003), and to a lesser extent by fixed-location gears, 

e.g. bottom longlines and bottom-set nets (Lumsden et al.

2007, Fuller et al. 2008). The 2007 report on the State of the

Deep Coral Ecosystems of the United States notes that use

of bottom-tending gears, especially bottom-trawls, is the

largest threat to deep-sea coral communities (Hourigan

et al. 2007). The report advocates further research on the

effects of fishing on deep-sea corals to inform management

decisions. This research priority was extended to sponges

in NOAA’s Strategic Plan for Deep-Sea Coral and Sponge

Ecosystems (NOAA 2010). Here, we briefly summarize

the known interactions between fishing gear and deep-sea

corals and sponges in U.S. waters.

Christopher N. 

Rooper1*, 

Peter J. Etnoyer2

, 

Kevin L. Stierhoff3, 

and John V. Olson4

• • •

1 NOAA Resource Assessment 

and Conservation Engineering Division, Alaska Fisheries 

Science Center, Seattle, WA

* Corresponding Author:

Chris.Rooper@noaa.gov

2 NOAA Center for Coastal 

Monitoring and Assessment,

National Centers for Coastal 

Ocean Science, Charleston, SC

3 NOAA Fisheries Resource 

Divison, Southwest Fisheries 

Science Center, Santa Cruz, 

CA

4 NOAA Habitat Conservation 

Division, Alaska Regional 

Office, Anchorage, AK
94

EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •

II. Recent Developments in

Research and Management

The effects of fishing gear on deep-sea corals 

and sponges are of considerable interest to 

fishermen, scientists, managers, environmental 

groups, and the general public. Since the 

2007 report on the State of the Deep Coral 

Ecosystems of the United States, there have 

been efforts to: 

1) Study the effects of mobile and fixedlocation fishing gears on deep-sea

corals and sponges;

2) Close areas where fishing may impact

deep-sea corals and sponges; and

3) Modify fishing gears and practices to

protect deep-sea corals and sponges.

Identifying areas where fishing overlaps 

the presence of deep-sea corals and sponges 

requires knowledge of fishing locations and 

deep-sea coral occurrences. Fishing locations 

are obtained from vessel monitoring systems 

(e.g., SAFMC 2010, 2013), logbooks or onboard observers. Coral and sponge occurrences 

are from observations and bycatch data for 

sponges and corals collected by the National 

Marine Fisheries Service (NMFS) observers 

(NMFS 2011, 2013) and bottom-trawl surveys 

(Clarke et al., this volume, Stone and Rooper, 

this volume). However, not all fisheries include 

observers, and not all observers collect bycatch 

data on corals and sponges (e.g. Packer et 

al., this volume). Yet bycatch data, anecdotal 

observations, and the spatial and depth overlap 

of fishing operations show that in all U.S. 

regions, some fishing is conducted in areas 

with deep-sea corals and sponges (Hourigan et 

al. 2007, NMFS 2011, NMFS 2013). 

Based on these types of bycatch data, 

models have been developed that predict the 

distribution of deep sea corals and sponges

(Bryan and Metaxas 2007, Sigler et al. 2015, 

Guinotte et al., this volume, Packer et al.,

this volume). These models can help identify 

areas of overlap of sensitive invertebrates

with fishing activity. However, many of

these models are for small areas, have coarse

spatial and taxonomic resolutions, and are

insufficiently informed by observation data.

II.1. Mobile Bottom-Contact

Fishing Gear

In U.S. waters, bottom-trawls are used on the 

continental shelf and upper slope, typically 

shallower than 500 m depth. In other regions of 

the world, bottom trawls are also used in deeper 

continental slope waters (Roberts 2002, Norse et 

al. 2012, Clark et al. 2016).

The effects of mobile bottom-contact fishing 

gear on deep-sea corals are well documented 

(reviewed by Clark and Koslow 2007, 

Hourigan et al. 2007, Clark et al. 2015), but less 

so for their effects on sponges (Van Dolah et 

al. 1987, Freese 2001, Wassenberg et al. 2002, 

Austin et al. 2007, Hogg et al. 2010). Impacts 

include damage and removal of the organisms 

(Freese 2001, Hall-Spencer et al. 2002, NRC 

2002, Reed et al. 2007), changes in community 

composition and productivity (Hiddink et al. 

2006, Hixon and Tissot 2007), and broad-scale 

modification of geology (Puig et al. 2012). 

For example, in the Aleutian Islands, damage 

to corals increased with fishing intensity 

(Heifetz et al. 2009), with the highest damage to 

antipatharians (15%), hydrocorals (9%) and 

gorgonians (6%) (Stone 2014). Additionally, 

coral removal by bottom-trawls accounted for 

over 92% of observed coral bycatch in west 

coast fisheries (Whitmire and Clarke 2007).

In the Gulf of Alaska, a single pass of a 

bottom-trawl removed 1,000 kg of Primnoa
95

coral and detached 27% of the colonies 

(Krieger 2001). Cumulative removals of coral 

by bottom-trawling in Alaska, from 1997 to 

1999, were estimated at 81.5 metric tons (mt) 

per year, totaling about 250 mt (NMFS 2004). 

More recently, the bycatch of corals (including 

sea pens and whips), bryozoans, and hydroids 

were reportedly 54 mt from 2003-2005 (NMFS 

2011) and 35.6 mt in 2010 (NMFS 2013). 

Comparable figures for Alaskan sponge 

bycatch were 238 mt in 2003-2005 and 271 mt 

in 2010 (NMFS 2011, 2013). 

NOAA has determined that year-round 

closures to mobile bottom-tending gear are 

particularly effective at protecting biogenic 

habitats comprised of deep-sea corals and 

sponges (NOAA 2010, Hourigan 2014). The 

world’s first bottom-trawl fishing closure, 

specifically designed to protect deep-sea 

corals, occurred in 1984 at Oculina Bank, a 

Habitat Area of Particular Concern (HAPC) 

off eastern Florida. In 1986, the entire 

Exclusive Economic Zone (EEZ) surrounding 

the U.S. Pacific Islands was closed to bottomtrawls and bottom-set nets and longlines. The 

Western Pacific Fishery Management Council 

had previously identified bottom-trawling by 

foreign fishing fleets as a threat to the region’s 

precious coral beds (Hourigan 2014).

Most other bottom-trawl closures have been 

enacted since 2005, primarily to protect EFH 

(Hourigan 2009, Hourigan 2014, Fig. 1). This 

includes bottom-trawl closures between 700 

and 1,750 ftm (1,280-3,200 m) on the west coast 

of the continental U.S., as well as 34 additional 

areas including biogenic habitats, banks, ridges, 

and canyons in March 2006. This management 

action protected 336,698 km2 (130,000 mi2) of 

marine habitat in Washington, Oregon, and 

California of EFH for groundfish.

In the Southeast U.S., five new areas 

containing Lophelia pertusa coral were closed

to bottom-contact fishing in 2010, and these

were expanded in 2015 to include newly

identified coral areas. In the same year, the

Mid-Atlantic Fishery Management Council

became the first council to use the discretionary 

provision of Magnuson-Stevens Act to propose

establishment of a deep-sea coral protection

area encompassing more than 38,000 mi2.

The Deep Sea Coral Amendment1

 includes

measures to restrict trawls, dredges, bottom

longlines, and traps within the bounds of the

protection area. Areas of the Aleutian Islands

in Alaska were closed to mobile bottomcontact fishing gear in 2006 (NMFS 2010). This 

decreased the bycatch of corals and sponges

under steady or decreasing fishing effort, until 

2011 (NPFMC 2012). However, data collected

through 2013 indicated an increase in bycatch 

of coral and sponges in the Aleutian Islands,

possibly due to increased fishing effort for 

rockfish in the region.

The cumulative effect of these management

actions between 2005 and 2010 was a sixfold increase in the total area of seafloor that

restricts bottom contact fishing gear within

the U.S. continental EEZ (Fig. 1). The majority

of the area closures in the U.S. occurred

in the Pacific and North Pacific Fishery

Management Council regions, and comprised

more than 1.35 million km2 (530,000 mi2) of

the U.S. EEZ. The cumulative area of these

closures represents 22% of the continental

US EEZ (2,401,232 mi2). For the most part,

these closures have not been monitored to

determine whether protections are effective in

recovering, maintaining or increasing deep-sea

coral diversity and abundance.

1 http://www.mafmc.org/actions/msb-am16

2 https://en.wikipedia.org/wiki/Exclusive_economic_zone

Modifications to mobile bottom-contact fishing

gear to reduce bycatch (e.g., turtle and marine

EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •
96

Figure 1. Cumulative areal extent of bottom closures by Fishery Management Council (FMC) region 

between 1982 to 2015 as a percent of EEZ in each region. Colors designate the closure type. US Regional 

FMCs are: GMFMC = Gulf of Mexico; SAFMC = South Atlantic; MAFMC = Mid-Atlantic; NEFMC = 

New England; NPFMC = North Pacific; PFMC = Pacific. The MAFMC closure (*) represents the deep-sea 

coral protection zones proposed in June 2015, covering 28,864 n mi2. The Western Pacific Council (WPFMC) and Caribbean Council (CFMC) are not shown. The WPFMC protected the entire EEZ (1,499,972 

nm2) under its jurisdiction from trawling and certain other bottom-contact fishing gears in 1983. Bottom-trawling does not occur in the CFMC region. The figure includes closures specifically designed to 

protect deep-sea corals and those using Essential Fish Habitat (EFH) authorities (NOAA 2011, Sutter et 

al. 2014). There are additional closures in place to reduce gear conflicts and other purposes, which may also 

protect deep-sea corals and sponges in such areas. Asterisk (*) indicates closures were proposed in 2015 and 

approved in 2016.

EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •
97

Figure 2. Commercial landings (millions of kg) of deep demersal species (> 50 m) in the Southern California Bight by gear type. Data are from California Department of Fish and Wildlife landings tickets (Perry 

et al. 2010; Perry et al. unpublished data) for select deep-water species codes during the time period 1972-

2011. 

 mammal excluder devices) and negative 

impacts on the seabed (e.g., reducing footrope 

or roller size on bottom-trawls) have been an 

active area of research within NMFS. Some of 

the gear modifications and restrictions have 

the potential to reduce impacts on deep-sea 

corals and sponges. For example, in Alaska, 

NMFS scientists have been working with 

flatfish bottom-trawl fisheries since 2002 to 

reduce bycatch and mortality of crab and 

other benthic invertebrates in soft substrates 

without negatively affecting catches of 

target species (Rose et al. 2010a, Rose et al. 

2010b, Rose et al. 2013). In those studies, the 

sweeps of bottom-trawls were raised off the 

seafloor using disk clusters spaced at 9-m 

intervals along the trawl sweeps. This gear 

modification reduced crab mortality, since 

the modified sweeps passed over many of the 

crabs it encountered, but had no significant 

effect on flatfish catch. Additionally, these 

gear modifications reduced, by almost half, 

the damage to sea pens under the path of the 

trawl gear (Rose et al. 2010a). Working closely 

with the fishing industry and the North Pacific 

Fishery Management Council, new regulations 

were implemented in 2011 that required 

bottom-trawl sweep modifications in the 

eastern Bering Sea flatfish fishery. On the U.S. 

west coast, where footrope size restrictions 

were instituted to reduce catches of rockfish 

species (Sebastes spp.), trawl fishing effort has 

EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •
98

shifted away from rugose habitats known to 

support corals and other sensitive benthic 

invertebrates (Bellman et al. 2005).

II.2. Fixed Bottom-Contact

Fishing Gear

In addition to mobile fishing gear, there are 

many types of fixed gear used in commercial 

fishing, including traps or pots, bottom-set 

nets, and longlines. Fixed gears are typically 

deployed in a wide range of depths, including 

depths > 1,000 m. Most of these gears are in 

contact with the seafloor when fishing, and 

are often used in areas, such as rocky reefs. 

These areas may be most important for deepsea corals and sponges that rely on rocky 

substrates for attachment.

In some areas, fixed-gears account for 

considerably more landings than mobile 

gears. For example, according to California 

Department of Fish and Wildlife (CDFW; 

formerly California Department of Fish and 

Game) annual landings reports (Perry et al. 

2010), the total landings by traps, set lines, and 

set nets in southern California can outweigh 

landings by bottom-trawls, especially in 

recent years (Fig. 2). Nearly 64% of recent 

(2007-2011) deep-water demersal landings 

from Southern California are from fixed gears 

(longlines and pots) for sablefish, thornyhead, 

hagfish, and spot prawn. The remaining 35% 

from mobile gear were predominantly soft 

sediment fisheries for sea cucumber and sole 

(Perry et al. 2010, Etnoyer et al. 2013). Fishing 

with fixed-gear (Fig. 3A) is often conducted 

further offshore, over a wider portion of 

the continental shelf and slope than mobile 

gear (Fig. 3B), yet probably covers a smaller 

cumulative area (e.g., NMFS 2005). 

Off southern California, and likely elsewhere, 

fixed-gear fisheries that target deep-water 

demersal species operate within the depth

ranges of deep-water corals and sponges with

the potential for negative interactions where

overlap occurs. Fixed fishing gears can interact

with corals in several ways. Pots and traps

can be dropped directly on top of colonies, or

dragged on the bottom during deployment

and recovery. Nets can snag on rocks, or

be cut loose during storms. Longlines may

become entangled during recovery, or when

hooked fish struggle. Incidentally, museum

specimens of the precious Corallium coral

were retrieved by longline fisheries surveys

off the Azores in 2005 (Sampaio et al. 2009). In

the U.S. Pacific Northwest, Paragorgia arborea

was toppled and wrapped in longlines in the

Olympic Coast National Marine Sanctuary

(Brancato et al. 2007) (Fig. 4A). Derelict fishing

gear has also been caught on coral in other

areas, such as Southern California (Fig. 4B-D),

Gulf of Mexico mesophotic reefs (Etnoyer et

al. 2015), Nova Scotia, Canada (Mortensen et

al. 2005), and in parts of the northeast Atlantic

Ocean (Sampaio et al. 2012).

There are, however, few peer-reviewed studies

that rigorously examine the effects of fixedgear fisheries on deep corals and sponges. A

recent study of small-scale, shallow

(< 25 m) artisanal fisheries in Mexico found

that bycatch rates were negligible for traps

and gillnets – less than 0.5 kg (0.12 kg of

gorgonians and 0.37 kg for sponges) for each

$1,000 of revenue generated by the fishery

catch – while bycatch rates of habitat-forming

invertebrates (as a percentage of total catch)

were ~20% for set gillnets and close to zero

for traps (Shester and Micheli 2011). Damage

and removal of 17% of shallow-water

gorgonian corals within a meter of a set net

was observed, while damage was minimal

and removals were zero for traps. Although

the impacted areas were relatively small for

these gear types, the cumulative impact may

EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •
99

Figure 3A. Distribution of demersal landings between 2007 and 2011 for deep-water fixed gear fisheries 

from California Department of Fish and Wildlife catch blocks in the southern California Bight (Perry et 

al. 2010; Perry et al. unpublished data). Also shown are the locations of gorgonian octocorals observed 

during remotely operated vehicle surveys since 2006 (NOAA 2015). 

EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •
100

Figure 3B. Distribution of demersal landings between 2007 and 2011 for deep-water bottom-trawls from 

California Department of Fish and Wildlife catch blocks in the southern California Bight (Perry et al. 

2010; Perry et al. unpublished data). Also shown are the locations of stony corals from surveys since 2006 

(NOAA 2015).

EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •
101

Figure 4. Images of longline and trap gear from remotely operated vehicle surveys on the US West Coast: 

A) Bubblegum coral (Paragorgia arborea) entangled in longline gear at ~200 m depth in Olympic Coast

National Marine Sanctuary (OCNMS; Brancato et al. 2007); B) Scleractinian coral (Lophelia pertusa)

colonies with yellow polypropylene line, possibly from an anchor line, or a buoy line on fixed gear;

C) trap gear (center) and netting (lower right) adjacent to a purple sea fan (Eugorgia rubens); and

D) a section of mesh from a trawl net wrapped around Lophelia pertusa rubble. Photo A is courtesy

NOAA OCNMS. Photos B-D are courtesy of NMFS, Southwest Fisheries Science Center.

be substantial given the large number of 

participants in the fisheries. In the Atlantic 

Ocean off Canada, a review of observer data 

indicated that the rate of coral occurrence in 

longline sets (13%) was higher than that of 

either gillnets (7%), trawls (4%), or crab pots 

(0%) (Edinger et al. 2007).

In the U.S., much of the data on bycatch 

rates for fixed gears comes from fisheryindependent surveys. For example, the catch 

rates for gorgonian corals in the Alaska 

longline survey that operates annually at 

stations on the slope of the eastern Bering Sea 

suggest very low rates of bycatch for corals 

and sponges (~2-4 coral colonies or sponges 

per station occupied by the survey, Fig. 5). 

Although some anecdotal evidence exists, 

there is generally less data available from 

U.S. fisheries to compare damage rates to 

coral and sponge habitat by fixed and mobile 

fishing gear. The most comprehensive study 

to date looked at damage rates in areas of 

EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •
102

the central Aleutian Islands that were only 

fished with fixed gear (pots and longlines) 

and found damage rates to be higher, but 

not significantly higher than damage rates in 

unfished areas (Heifetz et al. 2009). Removals 

of corals and sponges in Alaska by longline 

fisheries between 2003 and 2005 were 

estimated at 14 mt and 6 mt respectively, but 

these values constituted a minor fraction of the 

target species catch (NMFS 2011). Removals by 

pot fishing accounted for a negligible amount 

of coral bycatch and only 1 mt of sponge 

removal during the same time period. 

Removals of corals and sponges by fixed gears 

occur in other regions as well, but some of the 

evidence is anecdotal. For example, bamboo 

corals (Isidella sp.) have been reported in 

sablefish pots off California (S. Risherman, 

Marine Applied Research and Exploration, 

pers. comm.). Paragorgia bubblegum corals 

have been observed decorating the ceilings of 

salmon shacks in Neah Bay, Washington (P. 

Etnoyer, pers. obs.). Large black corals have 

been retrieved from trawl nets in Alaska, then 

polished and carved into decorative artifacts 

or jewelry (P. Etnoyer, pers. comm.).

In some places, derelict fishing gear may also 

damage deep-sea corals and sponges. During 

a rockfish study in the Aleutian Islands in 

2004, a juvenile rockfish was observed residing 

in a tangle of derelict longline gear (Rooper 

et al. 2007). During that study, 4.29 ha of 

seafloor was observed with an underwater 

video camera and three separate observations 

of derelict longline gear were observed among 

corals and sponges. 

Figure 5. Catch (mean number of colonies ± SE) of coral and sponge taxa in the Alaska Fisheries Science 

Center biennual longline survey of the eastern Bering Sea continental slope from 1997-2011. Data are 

available from the Alaska Fisheries Information Network (AKFIN, www.akfin.org).

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Observations from remotely operated vehicles 

(ROVs ) of derelict fixed gear in areas with 

deep-sea corals and sponges have also been 

documented in the northern Gulf of Mexico, 

where injuries to gorgonian octocorals were 

attributed to the snapper and grouper fisheries 

that comprise the reef-fish fishery (Etnoyer et 

al. 2015).

In the U.S., many areas have been closed to 

fishing to reduce the impacts of fixed gears 

on seabed habitats. For example, fifteen 

seamounts off the U.S. west coast and Alaska 

have been closed to all bottom-contact fishing 

gears since 2006, and a total of 7,020 m2 were 

restricted. Other examples include “Piggy 

Bank” and “The Footprint” essential fish 

habitat area in the Channel Islands National 

Marine Sanctuary, which harbors abundant 

coral and sponge species (Yoklavich et al. 2011, 

2013), that are now protected from bottom 

fishing of all types. Still, enforcement of these 

restrictions remains a challenge (C. Mobley, 

NOAA pers. comm.).

In a review of the impacts of gear on sensitive 

seabed habitats in the Southeast U.S., Barnett 

(2001) recommended excluding the use of 

bottom longlines in the vicinity of coral reefs 

due to their potential for entanglement. In this 

region, the use of bottom longlines, pots, and 

traps, as well as mobile gears is not allowed 

in the Oculina Banks HAPC, as well as in 

the larger deep-water Coral HAPCs, except 

in Golden Crab Allowable Fishing Areas or 

Shrimp Fishery Access Areas. We are not 

aware of any efforts to modify fixed bottomcontact gear to reduce their impacts. 

III. Rationale for Closures

It is important to recognize that most, if not 

all of the management actions and research 

on the interactions between fisheries and 

sensitive fauna have aimed to protect corals

and sponges while maintaining or enhancing

existing fisheries. In some cases, such as with

Oculina Banks and the Lophelia reefs in the

southeast U.S., the reef-like structure provided

by a single coral species has been the impetus

for protection measures. In others, such as

the “coral gardens” in Alaska, management

actions have focused on groups of taxa (such

as groups of corals and sponges separately)

rather than trying to protect an area for a

particular species. Corals and sponges have

also benefitted from research and management

actions that were driven by management

issues for other species. For example, trawl

modifications to reduce bycatch of overfished

species (as in the west coast footrope

restrictions example above) or non-target

species (as in the Alaska crab example above).

In the Gulf of Mexico, Lophelia reefs provide

habitat for golden crab (Chaceon fenneri),

which is fished using traps strung together

and recovered from the seafloor by grappling

hook. The deep-sea red crab (Chaceon

quinquedens) can aggregate in large colonies

of Madrepora oculata as deep as 1,000 m in the

Gulf of Mexico (Boland et al., this volume).

The fisheries for these species are not well

developed in the Gulf of Mexico, but they

do represent an emerging concern. The

South Atlantic Fishery Management Council

designated allowable fishing areas for golden

crab pot fisheries during the establishment of

Deepwater Coral HAPCs.

IV. Knowledge Gaps and

Future Challenges

In most regions of the U.S., methods have 

been developed and employed to study 

the effects or anticipated effects of mobile 

bottom-tending fishing gear. Most of this 

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104

knowledge has been gained in response to the 

EFH management mandate under the 1996 

amendment of the MSA, which emphasizes 

an ecosystem approach to management. In the 

last five years, the NMFS, in partnership with 

the fishing industry, has better characterized 

where fishing with bottom-tending gears is 

likely to occur. The extent that fishing overlaps 

with deep-sea coral and sponges is determined 

by the distribution of these organisms and 

fishing effort. For fisheries with observer 

programs, additional information on the 

locations and amounts of bycatch can provide 

some indication of the extent of interactions 

between fishing gear and these organisms. 

There is little information, however, on the 

relationship between bycatch observed on the 

vessel and the actual impact or encounter rates 

with corals and sponges on the seafloor. These 

impacts will vary among gear types, species 

assemblages, and seabed types. However, the 

effects of fixed gear on corals and sponges are 

poorly studied in most regions. 

A better understanding of the effects of fixed 

fishing gear on deep corals and sponges 

will come as more available data on the 

spatial extent contacted by this gear during 

fishing is revealed. With this information, 

the footprints of the various fixed gear types, 

and thus the area of the impacted seafloor 

can be calculated. There are also estimates of 

the bycatch of deep-sea corals and sponges 

from fishing gears from mandatory observer 

programs in Alaska (NMFS 2011). Still 

unknown is the performance of the gear 

during setting, fishing, and retrieval, and the 

amount of derelict gear that remains on the 

seafloor. We need to better understand the 

interactions between deep demersal target 

species, the gear types being used to fish them, 

and the types of corals and sponges occurring 

in fishing areas before we can develop optimal 

measures to protect these sensitive ecosystems 

while maintaining sustainable fisheries.

A second unknown is the degree to which the 

closed areas aid the recovery of damaged deep 

corals and sponges. Few studies quantify the 

densities, diversities, and distributions of deepsea corals and sponges before and after areas 

are closed. Althaus et al. (2009) studied stony 

coral habitats and associated megabenthic 

invertebrate assemblages on seamounts off 

Tasmania and found no clear signal of recovery 

after five years for habitats damaged by 

bottom-trawling; invertebrate communities 

remained impoverished, comprising fewer 

species at reduced densities than untrawled 

areas. Similarly, Williams et al. (2010) found 

no evidence of recovery in trawled seamount 

areas 5-10 years after protection in Australian 

and New Zealand. They concluded that the 

resilience of seamount ecosystems, dominated 

by corals, is low compared to most other 

marine systems disturbed by bottom-trawling. 

Studies to determine how quickly damaged 

ecosystems on the continental shelf and slope 

of the U.S. recover after a closure could inform 

optimal strategies for protecting areas and 

monitoring recoveries. 

V. Future Research and

Technology

Studies are needed that research pot and 

longline gear dynamics during deployment, 

fishing, and retrieval operations. These 

studies should be based on direct observation, 

remote sensing, or both. A number of new 

technologies may provide direct observations. 

Cameras on ROVs have been used to observe 

longline gear during fishing activities, but 

not during deployment or retrieval (K. L. 

Yamanaka, Department of Fisheries and 

Oceans, Canada, pers. comm.). Advances 

in camera technologies have also facilitated 

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105

observations. For example, researchers in 

the Australian Antarctic Division developed 

a camera system to collect video images 

periodically during longline deployments and 

retrievals (Kilpatrick et al. 2011). A modified 

version of the camera system is being 

developed at the Alaska Fisheries Science 

Center to deploy on commercial longline 

and pot gear. The intention is to collect 

stereoscopic images to measure objects and 

fish, and estimate the area that is contacted by 

the fishing gear during setting, deployment, 

and retrieval.

Multiple studies examined changes in coral 

and sponge communities in closed areas 

where trawling previously occurred (e.g. 

Freese 2001, Heifetz et al. 2009). In the Gulf 

of Alaska, there was no indication of sponge 

recovery after experimental trawling when 

the damaged sites were revisited a year later. 

These sites should be sampled on appropriate 

time and space scales to evaluate the recovery 

processes and periods. Following area closures 

off Oregon, the bycatch of coral appears to 

have declined for some fisheries (Fig. 6). 

However, it is unclear if bycatch reductions 

are due to area closures, gear restrictions, 

changes in abundances and distributions of 

coral and target species, or other factors that 

may influence fishing behavior (PFMC 2012). 

Few studies have examined differences in 

Figure 6. Coral bycatch (lb./km fished) from west coast trawl fisheries offshore Oregon state before (1 Jan 

2002 - 11 Jun 2006) and after (12 Jun 2006 - 31 Dec 2010) essential fish habitat closures (PFMC 2012).

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106

the community structure of deep-sea corals 

and sponges “before and after” area closures, 

or demonstrated how long and how many 

of these closures are needed to maintain the 

ecosystem services that deep-sea coral and 

sponge ecosystems provide.

VI. Conclusion

Since publication of the report on the State of 

Deep Coral Ecosystems of the United States 

in 2007, there have been a limited number 

of studies of the effects of fishing activity 

on deep-sea corals and sponges. In the U.S., 

most studies have focused on the effects of 

mobile fishing gear. However, studies are 

also needed on the likely, but perhaps lesser, 

effects of fixed-gears. Information from 

observer programs and scientific surveys 

have been instrumental in the identification 

of areas where high coral and sponge bycatch 

is occurring and for designating closed areas. 

Bycatch of non-target species has driven the 

development of fishing gear modifications, 

which has reduced their impacts on deep-sea 

corals and sponges.

VII. Acknowledgements

Thanks to Curt Whitmire, Janet Mason, Wayne 

Palsson, David Demer, and Gerard DiNardo 

for helpful reviews of this manuscript. 

Stephanie Nehasil conducted the analysis of 

CDFW landings data used to create Figure 

3. Thank you to Chris Lunsford and Cara

Rodgveller for providing data from the

AFSC longline survey, to Enrique Salgado

for assistance with Figures 2, 3, and 4; John

Butler and the SWFSC ROV team collected

the images used in Figure 4. Special thanks

to Catherine Polk for her gracious assistance

with the document layout.

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Sustainability Conference May 6-9, 2013. Pacific 

Fishery Management Council

Van Dolah RF, Wendt PH, Nicholson N (1987) 

Effects of a research trawl on a hard-bottom 

assemblage of sponges and corals. Fish Res 

5:39-54 

Wassenberg TJ, Dews G, Cook SD (2002) The 

impact of fish trawls on megabenthos (sponges) 

on the north-west shelf of Australia. Fish Res 

58:141-151

Whitmire CE, Clarke ME (2007) State of Deep Coral 

Ecosystems of the U.S. Pacific Coast: California 

to Washington. In: Lumsden SE, Hourigan TF, 

Bruckner AW, Dorr G (eds) The State of Deep 

Coral Ecosystems of the United States NOAA 

Technical Memorandum CRCP-3, Silver Spring, 

MD

Williams A, Schlacher TA, Rowden AA,

F, Clark MR, Bowden DA, Stewart R, Bax 

NJ, Consalvey M, Kloser RJ (2010) Seamount 

megabenthic assemblages fail to recover from 

trawling impacts. Mar Ecol 31:183-199

Woodby D, Carlile D, Hulbert L (2009) Predictive 

modeling of coral distribution in the Central 

Aleutian Islands, USA. Mar Ecol Prog Ser 

397:227-240

Yoklavich M, Laidig T, Krigsman L, Taylor A, 

Watters D, Love M, Lundsten L, Negrete, B 

(2011) A characterization of the coral and 

sponge community on Piggy Bank seamount 

in southern California from a survey using 

a remotely operated vehicle. Report to 

the NOAA Deep-Sea Coral Research and 

Technology Program, Silver Spring, MD. 63pp

Yoklavich M, Laidig T, Taylor A, Watters D, 

Krigsman L, Love M (2013) A characterization 

of the Christmas tree black coral (Antipathes 

dendrochristos) community on three seamounts 

in the Southern California Bight from a survey 

using a manned submersible. Report to NOAA 

Deep-Sea Coral Research and Technology 

Program, Silver Spring, MD. 82pp

EFFECTS OF FISHING GEAR ON DEEP-SEA CORALS AND SPONGES IN U.S. WATERS

• • •
         

       

   

Yellow Picasso sponge and white

sponges on Davidson Seamount.

Courtesy of NOAA/MBARI.
                       

     

                 

                   

                         

                 

                       

                   

                   

               

                 

                 

               

         

                     

                   

                 

                   

                 

               

                   

  

                   

                           

                   

             

     

   

     

   

   

     

   

   

     

   

     

   

     

     

       

   

     

         

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

STATE OF DEEP-SEA 

CORAL AND SPONGE 

ECOSYSTEMS OF THE 

U.S. WEST COAST 

M. Elizabeth

Clarke1*, Curt E. 

Whitmire2, and 

Mary M. 

Yoklavich3

  

1 NOAA Northwest

Fisheries Science Center,

Seattle, WA

* Corresponding Author:

Elizabeth.Clarke@noaa.gov

2 NOAA Fishery Resource

Analysis & Monitoring

Division, Northwest

Fisheries Science Center,

Monterey, CA

3 NOAA Fisheries Ecology

Division, Southwest

Fisheries Science Center,

Santa Cruz, CA

I. Introduction

The U.S. Pacific coast marine region encompasses the continental

margin off the coasts of Washington, Oregon, and California, and

accounts for about 7% (778,628 km2) of the total area of the U.S.

Exclusive Economic Zone (EEZ). The continental margin in this

region is characterized by a relatively narrow (5‐40 km) shelf and a

steep continental slope, with the shelf break at approximately 200

meters water depth. This area is wholly within the California

Current Large Marine Ecosystem (LME). The California Current

LME is a transition ecosystem between subtropical and subarctic

water masses and is characterized by seasonal upwelling. It

encompasses portions of two marine biogeographic provinces –

Oregon (cold‐temperate) and California (warm‐temperate [formerly

San Diego]) – with a boundary between Point Conception and Los

Angeles, CA (Allen and Smith 1988, McGowan 1971, Briggs and

Bowen 2012). Relatively large estuaries border the area, including

those associated with the Puget Sound, Columbia River, and San

Francisco Bay. Fishing for salmon, groundfish, small coastal pelagic

species, highly migratory species, and invertebrates (most notably

squid, crab, shrimp, and sea urchins) is significant throughout this

region.

Deep‐sea corals and sponges are found throughout the area (Austin

et al. 2007, Lee et al. 2007, Whitmire and Clarke 2007). Much of the

information on the general zoogeography of corals in the region

originated from taxonomic records collected from bottom trawl

Rockfish sheltering in a

glass sponge

(Aphrocallistes sp.) near

Grays Canyon off the coast

of Washington.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

surveys over the past century. Sea pens, black

corals, and gorgonians were recorded

coastwide in the catch of bottom trawl surveys

and by fishery observers monitoring bycatch

during commercial fishing operations. Our

understanding of the distribution of corals has

been informed by visual surveys using

underwater vehicles (e.g., submersibles,

remotely operated vehicles [ROVs] and

autonomous underwater vehicles [AUVs]).

Whitmire and Clarke (2007) reported 101

species of corals from six cnidarian orders from

the West Coast region. They highlighted

significant coral communities at Davidson

Seamount (DeVogelaere et al. 2005), Monterey

Canyon, Cordell Bank (Pirtle 2005), the

Olympic Coast National Marine Sanctuary

(Hyland et al. 2005, Brancato et al. 2007), and on

numerous rocky banks off southern California

(Yoklavich and Love 2005, Tissot et al. 2006,

Love et al. 2007).

Major stressors on deep‐sea coral communities

off the Pacific Coast included oil and gas

development in the Southern California Bight,

deployment of pipelines and communication

cables, and marine pollution. Fishing

operations, particularly bottom trawling, were

identified as the most immediate and

widespread threat to these communities. The

risk posed by fishing activities was reduced by

fishery management measures (e.g., area

closures such as marine protected areas (MPAs)

and gear restrictions) taken before 2007. In June

2006, the Pacific Fishery Management Council

(PFMC), in cooperation with NMFS,

implemented a comprehensive plan to protect

essential fish habitat (EFH) for west coast

groundfish (PFMC 2005, 2006). This plan

identified habitat areas of particular concern

(HAPCs) that protect vulnerable biogenic

habitats. The plan was developed in

collaboration with non‐governmental

organizations, the fishing industry, and the

National Marine Sanctuary Program, and

addressed impacts from a broad range of

human activities (fishing and non‐fishing) and

included procedures for review as new

information became available.

As of 2007, MPAs were distributed along the

length of the Pacific coast in both federal and

state waters. Over 130,000 mi2 (336,700 km2)

had been designated as MPAs fully protected

from impacts from bottom trawls, with selected

vulnerable habitats protected from all fishing

gears that contact the bottom. Additional MPAs

and fishery closures were designated by the

individual states, including bottom trawl

prohibitions for the entire portion of

Washington state waters and most of California

state waters. Furthermore, there are five

national marine sanctuaries in the region, all of

which prohibit 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 this report, we highlight new scientific

information on deep‐sea corals uncovered since

the 2007 report, and outline new management

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

measures that help protect these organisms on

the Pacific coast. We also include new

information on deep‐sea sponges, which were

not previously addressed in the 2007 report.  

II. Update on Scientific

Information

II.1. New Research – Overview

Several new studies on both deep‐sea corals

and sponges were initiated or completed since

2007 by researchers from NOAA and a variety

of collaborative institutions. This includes

research funded by NOAA’s Deep Sea Coral

Research and Technology Program, other

targeted studies conducted by academic and

non‐governmental institutions, and work

aboard ships of opportunity, all of which has

yielded valuable information about corals,

sponges and their habitats. The program also

supported analysis of archived video or

photographic surveys to assess the occurrence

of deep‐sea coral and sponge habitats. There

also has been an expansion of regional

bathymetric mapping of the seafloor (PFMC

2012), and of new methods to predict suitable

habitats for corals. In addition, considerable

attention has been given to the genetics of

corals, which will assist in their identification

and taxonomy. Opportunistic data collections

continue, with coral and sponge specimens

taken in the annual west coast groundfish

bottom trawl survey and by observers

monitoring the groundfish fishery.  

New field research on deep‐sea coral and

sponge communities has been conducted coast‐

wide, with many study sites located in and

around the five national marine sanctuaries

(Figure 1; Appendix Table A). Additional field

expeditions, while not specifically targeting

corals or sponges, also have provided

information on these taxa.

II.1.i – NOAA’s Deep Sea Coral Research

and Technology Program

In 2010, NOAA’s Deep Sea Coral Research and

Technology Program launched a research

initiative on the U.S. West Coast to locate,

study, and provide information for the

conservation of deep‐sea coral and sponge

habitats. The initiative was developed in

consultation with the PFMC and targeted

information that could inform management.

Field research was conducted at various sites in

and around four national marine sanctuaries

and in sponge habitats offshore of Grays

Harbor, Washington as well as at seamounts in

Canadian and international waters to further

the understanding of biogenic habitats and

associated fishes (Figure 1). These areas were

chosen based on expected suitability for deep‐

sea coral or sponge habitats and their potential

to inform sanctuary management plans and the

PFMC’s five‐year review of groundfish EFH.

The field research, originally planned for three

years (2010‐2012), was extended to 2014 and has

made substantial progress.

During this West Coast initiative, field research

addressed one or more of the following specific

goals:

1. Locate and characterize coral and sponge

habitats, including the following:

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

a. Collection of baseline data on

abundance, size, condition, and

distribution of deep‐sea corals and

sponges;

b. Mapping of targeted seabed and

collection of environmental data

associated with coral and sponge

habitats (e.g., depth, sea floor

substratum types, seawater

temperature, salinity, pH, and dissolved

oxygen) to help understand habitat

factors that may influence species

distribution.

2. Quantify fish and invertebrate associations

with corals and sponges to help understand

their potential value as habitat.

3. Assess the condition of coral and sponge

assemblages in relation to potential

anthropogenic or environmental

disturbances.

4. Collect specimens of deep‐sea corals,

sponges, and associated organisms to

confirm taxonomic identifications, and for

genetic and reproductive analyses.

Program‐funded surveys were conducted using

remotely operated vehicles (ROVs), human

occupied vehicles (HOVs), and an autonomous

underwater vehicle (AUV). Observations of

corals and sponges are being incorporated into

NOAA’s National Database for Deep‐Sea

Corals and Sponges.

Figure 1. General locations of surveys of deep‐sea

corals and sponges off the Pacific coast from 2007 to

2013, including NOAA’s Deep Sea Coral Research

and Technology Program initiative. See Appendix

Table A for details of individual surveys.

II.1.ii – Additional field research

Researchers at a number of institutions

conducted studies that contribute to our

knowledge of deep‐sea corals off Washington,

Oregon and California (Appendix Table A). Of

particular significance are deeper water surveys

by the Monterey Bay Aquarium Research

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Table 1. Summary of deep‐sea coral and sponge taxa recorded during bottom trawl tows as part of the West

Coast Groundfish Bottom Trawl Survey (2001‐2012). “n” denotes number of tows with recorded bycatch;

“FREQ” denotes ratio of tows with catch to total tows recorded; “Weight” denotes catch weight (kg); “FREQ

Rank” and “Weight Rank” columns denote the rank of taxon by either frequency of occurrence or catch

weight, respectively. Taxa are ranked by highest catch weight within major taxonomic groups.

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Taxon FREQ Weight Weight Taxon n FREQ Rank Rank (kg) Rank

Porifera Phylum 1,436 19.9% 1 11,035.01 1

Hexactinosida Order 512 7.1% 3 7,992.77 2

Rossellinae Subfamily 159 2.2% 8 1,328.24 3

Suberites spp. Genus 20 0.3% 19 1,073.27 4

Hyalonema spp. Genus 180 2.5% 7 561.63 5

Hexactinellida Class 17 0.2% 21 131.01 6

Farrea spp. Genus 11 0.2% 25 4.18 24

Porifera 1,869 25.9% 22,126.11

Pennatulacea Order 926 12.8% 2 102.11 7

Anthoptilum grandiflorum Species 489 6.8% 4 97.89 8

Halipteris spp. Genus 211 2.9% 5 29.68 12

Ptilosarcus gurneyi Species 135 1.9% 10 21.09 14

Umbellula spp. Genus 141 2.0% 9 8.84 18

Anthoptilum murrayi Species 42 0.6% 15 3.00 25

Anthoptilum spp. Genus 5 0.1% 27 0.36 30

Pennatula phosphorea Species 15 0.2% 22 0.28 32

Halipteris willemoesi Species 5 0.1% 27 0.14 33

Halipteris californica Species 3 0.0% 28 0.05 34

Pennatulacea 1,670 23.1% 263.44

Gorgonacea Order 197 2.7% 6 72.55 9

Paragorgia spp. Genus 26 0.4% 18 11.69 17

Isidella spp. Genus 12 0.2% 24 7.15 19

Callogorgia kinoshitae Species 29 0.4% 17 6.31 20

Keratoisis spp. Genus 5 0.1% 27 1.95 26

Callogorgia spp. Genus 5 0.1% 27 0.33 31

Leptogorgia chilensis Species 1 0.0% 30 0.04 35

Swiftia pacifica Species 2 0.0% 29 0.02 37

Acanthogorgiidae Family 1 0.0% 30 0.01 38

“Gorgonacea” 253 3.5% 100.04

Antipatharia Order 126 1.7% 11 64.26 10

Chrysopathes spp. Genus 46 0.6% 13 44.45 11

Bathypathes spp. Genus 44 0.6% 14 1.37 27

Lillipathes spp. Genus 18 0.2% 20 1.01 28

Antipatharia 217 3.0% 111.09

Heteropolypus spp. Species 104 1.4% 12 21.93 13

Anthomastus spp. Genus 39 0.5% 16 11.70 16

Alcyonacea Order 29 0.4% 17 6.11 21

Nephtheidae Family 1 0.0% 30 0.04 36

Alcyonacea 168 2.3% 39.78

Caryophylliidae Family 7 0.1% 26 12.60 15

Flabellidae Family 13 0.2% 23 5.86 22

Scleractinia Order 7 0.1% 26 5.34 23

Scleractinia 26 0.4% 23.80

Stylasteridae Stylasteridae Family 5 0.1% 27 0.53 29

ALL TAXA 3,182 44.0% 22,664.78
                       

     

                

           

             

           

               

             

                 

           

         

       

       

             

           

               

      

             

               

                 

           

               

             

           

                 

       

           

       

             

         

           

             

           

           

               

           

           

             

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Institute on a number of seamounts with rich

deep‐sea coral and sponge habitats. Additional

field research has been conducted in the

national marine sanctuaries (e.g., Etherington et

al. 2011), by Oceana (Shester et al. 2012,

Enticknap et al. 2013), as part of monitoring

state MPAs (e.g., Yoklavich et al. 2010), and in

association with other studies. Several graduate

students at Washington State University

summarized information on the spatial

distribution and abundance of invertebrates

and fishes that were quantified from archived

videotape collected during visual surveys using

ROVs or a submersible (e.g., Bright 2007, Graiff

2008, Bianchi 2011).

II.1.iii – Bottom trawl collections

The occurrence of deep‐sea corals and sponges

as catch in the NMFS West Coast Groundfish

Bottom Trawl Survey remains the most

comprehensive source of coast‐wide

information on the general distribution of

higher‐level coral and sponge taxa. These

surveys are conducted annually in “trawlable”

habitats coast‐wide at depths from 55 to 1280

m. From 2001‐12, sponges and pennatulaceans

were the most abundant taxa observed in these

surveys (Table 1), most likely because of the

types of habitats (e.g., low relief) that are

accessed by bottom trawls. Physical specimens

were collected, with some sent to taxonomists

for identification.  

These data were summarized for the five‐year

review of Pacific Groundfish EFH conducted by

NMFS and PFMC (PFMC 2012). This analysis

focused on changes in bycatch before and after

implementation of EFH regulatory measures in

June 2006. Although no significant difference in

the total frequency of occurrence of corals and

sponges is apparent, identification of species in

the trawl surveys is improving and often

samples are retained and sent to experts for

taxonomic verification of species. Information

collected by observers of the groundfish fishery

provides only broad taxonomic resolution;

however, these observers routinely collect

specimens for species verification.

II.2. Taxonomy and Species

Distributions

II.2.i – Corals

a. Coral taxonomy

Whitmire and Clarke (2007) reported 106 taxa

of corals within the U.S. EEZ from the West

Coast region (Table 2). The list included 52 taxa

with incomplete taxonomy. Since 2007, there

have been several additions to the number of

deep‐sea coral species documented on the west

coast including range extensions of certain

species (Tables 2 and 3; Whitmire et al. 2017,

Online Annex). In addition, the relationships

between species have been reviewed and

several species have been synonymized.

With funding from the Deep Sea Coral

Research and Technology Program, samples

(vouchers) of deep‐sea corals from the

northeast Pacific Ocean have been identified to

species using both morphology (form and

structure) and DNA sequence bar codes

(Everett et al. 2016). Coral samples have been

collected during field research, research trawl

surveys and by fishery observers. Genetic

sequences for two mitochondrial genes – COI

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Table 2. Number of coral taxa reported from the U.S. West Coast region in 2007 and in the present report.

Increases include new species identifications and range extensions (Table 3), as well as species not included in

the first report. Data sources and references online in Whitmire et al. 2017.

.TAXON Whitmire &

Clarke 2007

Whitmire et

al. 2017

Class Anthozoa

Black corals (Order Antipatharia) 8 9

Stony corals (Order Scleractinia) 18 19

Gorgonian corals (Order Alcyonacea – in part

[formerly Order Gorgonacea]) 39 60

True soft corals and stoloniferans (Order

Alcyonacea) 8 13

Sea pens (Order Pennatulacea) 28 28

Hydrozoa

Stylasterid corals (Order Anthoathecata, Family

Stylasteridae) 5 6

Total 106 135

and MutS – are linked to voucher specimens to  

create a resource for researchers of deep‐sea

corals. Microsatellite loci were also developed

for the gorgonian Primnoa pacifica Morrison et

al. 2015) and a species of sea pen in order to

analyze genetic patterns in populations of deep‐

sea octocorals. By creating a voucher collection

for deep‐sea corals and linking those vouchers

to species‐specific genetic sequences,

subsequent taxonomic identifications can

potentially be made using DNA sequences

rather than sending individual specimens to the

few morphological taxonomists (Everett et al.

2017). The genetic information also will be

useful in the identification of corals collected

from fishery observers and bottom trawl  

surveys, where poor condition of specimens

makes morphological identifications difficult.  

b. Coral distribution

Based on recent studies, an improved

understanding of the distribution of coral

species is emerging on the West Coast. With the

exception of pennatulaceans, corals mostly

occur on rocky substrata (e.g., boulders,

pinnacles, rock outcrops). Region‐wide, hard

and mixed seabed types are relatively rare,

representing only around 10% of the substrata

from the shelf to depths of 3000 m (NMFS

2013). These seabed types are often associated

with seamounts, banks, and canyons, which

have been the focus of many recent studies.

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Table 3. New coral species and range extensions reported from the U.S. West Coast region since 2007.

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Order Species Reference

New species:

Gorgonians Chrysogorgia pinnata

Chrysogorgia monticola

Calyptrophora bayeri

Calyptrophora laevispinosa

Narella alaskensis

Cairns 2007

Cairns 2007

Cairns 2007

Cairns 2007

Cairns and Baco 2007

Narella bowersi Cairns 2007

Parastenella gymnogaster

Parastenella pacifica

Isidella tentaculum

Sibogagorgia cauliflora

Leptogorgia filicrispa

Swiftia farallonesica  

Cairns 2007

Cairns 2007

Etnoyer 2008

Herrera et al. 2010

Horvath 2011

Williams and Breedy, 2016

Soft Corals Gersemia juliepackardae Williams and Lundsten 2009

Synonymized

Gorgonians

species:

Euplexaura marki = Chromoplexaura

marki

Williams 2013

Soft Corals Gersemia rubiformis = Alcyonium sp. Williams 2013

Range Extensions:

Scleractinians

Gorgonians

Pennatulaceans

Caryophyllia diomedeae

Caryophyllia quadragenaria

Paragorgia arborea (CA Province)

Paragorgia stephencairnsi (CA Province)

Parastenella ramosa (CA Province)

Swiftia pacifica (CA Province)

Eugorgia rubens (OR Province)

Pennatula phosphorea (CA Province)

Umbellula lindahli (OR Province)

Gonzalez‐Romero et al.

Gonzalez‐Romero et al.

Yoklavich et al. 2011

Yoklavich et al. 2011

Yoklavich et al. 2011

Yoklavich et al. 2011

M. Yoklavich (unpubl.)

Yoklavich et al. 2011

M. Yoklavich (unpubl.)

2009

2009

Lundsten et al. (2009) described the abundance

and distribution of benthic megafaunal

invertebrates on Pioneer, Davidson, and

Rodriquez seamounts off central and southern

California. Coral diversity was relatively high

with 23, 25, and 26 species identified,

respectively on these seamounts. Davidson

Seamount was the site for several detailed

studies of community structure and endemicity

(McClain et al. 2009, McClain et al. 2010). From

these studies, there was little support for

seamount endemicity, but seamount

communities may have unique structure that

changes significantly with depth. The summit

of Davidson Seamount (1299‐1477 m) was

characterized by the large bubblegum coral

Paragorgia arborea and several hexactinellid

sponge species (Farrea occa, Heterochone calyx,

and Chonelasma sp.), while deeper zones (> 1600

m) were characterized by different corals (e.g.,

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Corallium sp., Chrysogorgia spp., bamboo corals

and black corals) and other abundant glass

sponge species (McClain et al. 2010). Dense

aggregations of large corals occurred on peaks

and steep slopes, but were less abundant in

valleys or on the flanks (Clague et al. 2010).

Deep‐sea coral records from Monterey Bay

Aquarium Research Institute ROV surveys at

San Juan and San Marco seamounts also

revealed relatively high densities of gorgonian

and antipatharian taxa that commonly reach

large sizes (e.g., Paragorgia spp., certain isidid

bamboo corals, and certain black corals).  

In 2011 and 2012, two surveys were conducted

on Canada’s Bowie Seamount and on Cobb

Seamount just outside the U.S. EEZ, by the

Canadian Department of Fisheries and Oceans

in collaboration with NMFS researchers.

Information particularly on coral species

occurring on Cobb Seamount was expanded

during these cruises: 8 of the 11 species of

gorgonians and soft corals (Order Alcyonacea)

and all of the species of black corals (Order

Antipatharia) were new records for Cobb

Seamount (Curtis et al. 2015, Du Preez 2015).

In addition to seamounts, the Southern

California Bight has varied topography with

rocky banks, island flanks and ridges, and

submarine canyons. A survey on Farnsworth

Bank on the seaward side of Catalina Island

revealed that much of the rocky area is covered

by the purple hydrocoral, Stylaster californicus,

at depths to 66 m (Love et al. 2010). Bright

(2007) analyzed video from surveys conducted

with a manned submersible in 1995‐2004 at

depths of 97‐314 meters at the “Footprint,” a

high‐relief rocky bank south of Anacapa Island

in the Channel Islands. Densities of corals > 20

cm in size were quantified. Gorgonians were

relatively common (7/100 m2) with a mean size

of 30 cm. Black corals, predominantly

Antipathes dendrochristos, also were relatively

abundant (4/100 m2). These densities were

higher than reported by Tissot et al. (2006) for

other banks in the region but much lower than

more recent studies conducted in this area.

In 2010, deep‐sea corals and sponges, along

with fishes and seafloor habitats, were

surveyed using the Kraken II ROV and the

Seabed AUV at depths of 275‐900 m on Piggy

Bank near the Footprint Bank within the

Channel Islands National Marine Sanctuary. At

least 26 taxa each of corals and sponges were

quantified (Yoklavich et al. 2011). This site

comprised some of the highest densities of

corals and sponges found in the West Coast

region, ranging from 10‐54 corals/100 m2 and

22‐97 sponges/100 m2. The most abundant

corals (depending on depth) included

Christmas tree black coral (A. dendrochristos),

mushroom soft coral (Heteropolypus ritteri),

several species of Primnoidae and Plexauridae

(Swiftia spp.), dense stands of Lophelia pertusa

and cup corals (Desmophyllum dianthus), and the

sea pen (Halipteris californica; only on soft

sediment).  

Visual surveys using a manned submersible in

2010 focused specifically on the biology and

ecology of Christmas tree black corals on deep

offshore banks within the Channel Islands

National Marine Sanctuary (Yoklavich et al.

2013). This survey included habitats of high‐

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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relief rock boulders and outcrops and steep

slopes of soft sediments and rock rubble at

depths 110‐475 m. The 272 Christmas tree corals

observed during 11 dives were 5‐200 cm in

height, and most were healthy with little or no

evidence of damage.  

Over 23,500 deep‐sea corals and 23,800 sponges

also have been identified from archived video

transects conducted from a manned

submersible while surveying fishes on rocky

banks off southern California from 1995 to 2008

(M. Yoklavich, NOAA Deep‐Sea Coral and

Sponge Database). Various types of sea fans

were among the most abundant corals,

including golden corals (Acanthogorgia spp.),

primnoids (Plumarella longispina), and

plexaurids (Swiftia spp.). scleractinian cup

corals and substantial colonies of Lophelia

pertusa also were relatively abundant. Sponges

from at least 18 taxa in a variety of shapes,

sizes, and colors were observed. These data

have been integrated into NOAA’s National

Database for Deep‐Sea Corals and Sponges

(https://deepseacoraldata.noaa.gov/).  

A photographic database of 30,000+ images

collected during ROV surveys 2003‐2011 in the

Southern California Bight recently has been

reviewed (unpubl. data, P. Etnoyer and K.

Stierhoff). Distinct depth‐related patterns were

observed in coral assemblage structure. Shallow

depths (< 100 m) were dominated by Leptogorgia

chilensis, Stylaster californicus, and Eugorgia

rubens, while between 100 – 400 m

Plumarella sp. Lophelia pertusa, Acanthogorgia sp.,

Antipathes sp., and Paragorgia sp. were the most

abundant taxa.  

Deep‐sea corals also are associated with banks

off Central California (e.g., Cordell Bank: Pirtle

2005, Etherington et al. 2011; Rittenburg Bank:

Etnoyer et al. 2014) and Oregon (e.g., Coquille

Bank: Strom 2006, Stierhoff et al. 2011,

Enticknap et al. 2013). At Coquille Bank, the

most abundant gorgonian was the small red

Chromoplexaura marki (Stierhoff et al. 2011,

Williams 2013). At least one specimen was

observed with catshark egg cases attached.  

Deep‐sea corals are often observed on hard

substrata in canyons. Bianchi (2011) assessed

invertebrate composition, habitat complexity,

and ecological associations with structure‐

forming invertebrates and fishes in three

submarine canyons: Astoria Canyon off

northern Oregon, and Carmel and Ascension

canyons off the central California coast. There

were two distinct habitat assemblages

associated with soft and hard substrata in all

three canyons, and deep‐sea corals were mostly

small species at relatively low densities (0.4‐5

per 100 m2). In 2011, a survey was conducted

using an AUV to assess the distribution of

corals and sponges in Bodega Canyon, an area

under consideration for possible expansion of

the Cordell Bank National Marine Sanctuary.

Much of this canyon had been categorized as

hard substrata based on earlier multibeam

sonar mapping. However, from the visual

survey using the AUV, this area primarily was

found to be mud draped rock with only

occasional rocky outcrops and few small corals

and sponges (Fruh et al. 2013).

Several studies have reported information on

deep‐sea coral and sponge habitats and

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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Figure 2. Corals and sponges of the West Coast: a) Deep‐sea corals off Northern California at 750 m depth,

including Swiftia sp., Paragorgia sp., Parastenella sp., and a hexactinellid sponge. b) Habitat dominated by

deepwater sponges at Rittenburg Bank, Gulf of the Farallones National Marine Sanctuary. c) Fishing gear line

wrapped around Lophelia pertusa coral on Seamount 109 in the Channel Islands. d) Heterochone calyx glass

sponge from the Channel Islands National Marine Sanctuary. e) Small colonies of Swiftia sp. and Lophelia

pertusa on Piggy Bank off Southern California. f) Stylasterid corals on rocky banks in the Channel Islands.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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demersal fishes within Monterey Bay National

Marine Sanctuary, including Graiff (2008), Starr

and Yoklavich (2008), Yoklavich and Starr

(2010), Bianchi (2011), Knight (2011), Stierhoff et

al. (2011), and Shester et al. (2012). Most of

these studies were conducted in depths of 20‐

365 m using visual survey techniques from a

manned submersible and ROVs. Scientists

conducted baseline monitoring of seafloor

communities in and out of eight newly

                                   

                       

                               

                                 

                                       

                                       

                              

Figure 3. Distribution of coral (green) and sponge (orange) catch weights (kg) for tows conducted as part of

the Northwest Fisheries Science Centerʹs annual West Coast Groundfish Bottom Trawl Survey (2001‐2012).

Symbol sizes are proportional to size of catch for all coral (excluding pennatulaceans) and sponge taxa

recorded in the trawl net. Maximum sponge catches per tow within the time period were significantly higher

than coral catches, so circle size is equal to radius (km) for corals and area (km2) for sponges. The survey

extent (irregular polygon) is bounded by the exclusive economic zone to the north and south and by the 30 and

700 fathom isobaths. Tows with no catch of either corals or sponges are not shown.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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designated MPAs on the central California in

2007 and 2008 (Yoklavich and Starr 2010).

Direct observations were made of 134,000 fishes

and 42,098 structure‐forming invertebrates in

341,000 m2 of seafloor habitats that included

deep rocky banks and outcrops, canyons,

cobble fields and mud flats. At least 16 taxa of

corals and 14 taxa of sponges have been

cataloged; abundant corals were Stylaster

californicus, Leptogorgia chilensis, and

Heteropolypus ritteri. Lophelia pertusa was

relatively common in rocky steep areas. ROV

surveys on Rittenburg Bank, Farallon

Escarpment, and Fanny Shoals in the Gulf of

the Farallones National Marine Sanctuary

revealed twenty taxa of corals and sponges

(GFNMS 2013, Etnoyer et al. 2014).

Surveys of corals and sponges in Olympic

Coast National Marine Sanctuary have received

significant attention (Brancato et al. 2007;

Bowlby et al. 2011a, 2011b). While the dominant

corals in many areas were smaller species (e.g.,

Swiftia spp., Calcigorgia beringi, and Stylaster

sp.), aggregations of larger species (Paragorgia

arborea and Primnoa pacifica) also occurred in

several locations.  

The NMFS West Coast Groundfish Bottom

Trawl Survey provides a broader view of deep‐

sea coral distributions at fishable depths (i.e.,

55‐1280 m) along the whole West Coast (Keller

2017). In these surveys, most corals were

identified to order. The largest catches of corals

by weight occurred in 800‐1200 m of water near

the Oregon‐California border (Figure 3). Taxa

in these high‐weight catches included

gorgonian, black, and cup corals. This area also

exhibited some areas of high bycatch of corals

in the commercial trawl fishery as documented

by the West Coast Groundfish Observer

Program (PFMC 2012).  

Several of these high‐bycatch areas were

subsequently surveyed by ROV, towed camera,

and AUV, revealing both high densities of

deep‐sea corals, as well as evidence of damage

from fishing gear (Yoklavich et al. 2016,

Yoklavich et al. 2017).

c. Modeling distribution and abundance

of corals

Modeling efforts relevant to the distribution

and abundance of deep‐sea corals on the West

Coast are less than 10 years old, but the field is

rapidly advancing. Five modeling efforts that

incorporated either presence only or presence–

absence data of various coral taxa and

associated environmental factors off the west

coast have been conducted recently. Bryan and

Metaxas (2007) modeled habitat suitability for

two gorgonian families (Primnoidae and

Paragorgiidae) in the Northeast Pacific from

northern California to Alaska. Guinotte and

Davies (2012, 2014) used presence‐only data

and a variety of environmental factors to

develop predictive models of habitat suitability

for several broad taxonomic groups of deep‐sea

corals for the entire West Coast region (see

Guinotte et al., this volume). Slope,

temperature, salinity, and depth were

important predictors for most taxa in their

study. The majority of predicted suitable

habitat for most taxa occurred within areas

protected from bottom‐trawling (e.g., 82% for

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a. Sponge taxonomy

             

                 

                 

               

         

               

           

             

             

   

               

               

             

             

             

               

             

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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black corals and 66% for calcaxonian

gorgonians); however, the majority of suitable

habitat for holaxonian gorgonians (64%) was

predicted to occur in unprotected areas. Both

region‐wide studies lacked high‐resolution

bathymetric and substratum data.

Deep‐sea coral habitat modeling also has been

conducted from data collected on Cordell Bank.

Predictive models using presence‐absence data

and generalized linear models of Stylaster spp.

and Swiftia spp. indicated that small‐scale

habitat features, including bathymetry, slope,

and topographic position index, were

important habitat predictors for both taxa

(Etherington et al. 2011). Bathymetry and

topographic position index had a positive

influence, while slope had a negative influence

on these taxa. Rugosity had a strong influence

on the presence of Stylaster spp., which were

associated with shallow, rocky habitats with

high slope. Substratum type and aspect

significantly contributed to the presence of

Swiftia spp., which were predicted to be

broadly distributed in deep water over a

diversity of substratum types with low slope.

Krigsman et al. (2012) developed generalized

linear models to predict the probability of

occurrence of five commonly observed taxa

(stony cup corals, hydroids, short and tall sea

pens, and brittle stars) off southern California

using presence‐absence data from video

transects in the Santa Barbara Channel.

Predictive maps of probability of occurrence

were produced using validated models and

covariates of depth, location, and seafloor

sediment types. Bottom habitat type was the

strongest predictor of the occurrence of stony

cup corals, which occurred predominantly on

mixed sediment and rugose rock. As expected,

sea pens were predicted to occur on

unconsolidated sediments, but high

probabilities of occurrence were predicted only

in areas where bottom trawling was relatively

low.

Most recently, models have been developed to

predict the densities of the black coral (A.

dendrochristos) using environmental factors such

as depth, ocean currents, and productivity

throughout the Southern California Bight (Huff

et al. 2013). Predicted densities have been

mapped in relation to the extensive system of

MPAs throughout the Bight.

II.2.ii – Sponges

There have been few systematic studies of

sponge taxa for the U.S. West Coast. Austin et

al. (2007) produced a list of sponges for the

northeast Pacific, and Lee et al. (2007) compiled

current information on the sponges of

California. Many of the species from the region

were described from collections conducted in

the late 1800s and early to mid‐1900s.

From surveys off southern California, a new

species of carnivorous demosponge, Cladorhiza

pteron, was collected from 1,442 m depth on San

Juan Seamount, and also was seen at Rodriguez

Seamount (Reiswig and Lee 2007). Lundsten et

al. (2014) described three additional species of

cladorhizid sponges off the west coast. Several

of 13 sponges collected during an ROV survey

on Piggy Bank off southern California were

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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new records for the northeastern Pacific and

likely new species (pers. comm., W. Austin,

Khoyatan Marine Laboratory, Sidney, BC;

Yoklavich et al. 2011). In addition to new

records and new species that have been

reported since the 2007 report, several recently

published taxonomic guides and compilations

have improved the identification of sponges

and corals on the West Coast and Alaska

(Burton and Lundsten 2008, Stone et al. 2011).

b. Sponge distribution

Sponges occurred in about one quarter of all

trawls by the NMFS West Coast Groundfish

Bottom Trawl Survey (Table 1), though most

could only be identified as “Porifera.”

Compared to corals, the focus on identifying

sponges is very recent. The identified sponges

were predominantly glass sponges (Class

Hexactinellida) and demosponges in the genus

Suberites. The highest weight of sponges per

tow occurred in relatively discrete areas,

especially off the coast of northern Oregon and

around the northern Channel Islands, CA

(Figure 3).

In general, taxonomic information on sponges

is lacking, and species identification of sponges

from visual surveys is even more difficult than

for corals. Consequently, much of the

distributional information is based on

morphotypes (e.g., vase, shelf, foliose). Sponges

have been reported from nearly all the surveys

in rocky habitats off the West Coast that

reported deep‐sea corals. In many cases

sponges were larger and more abundant than

corals (other than sea pens). Bianchi (2011)

reported higher densities and larger sizes for

sponges than for corals in Astoria, Ascension

and Carmel Canyons. Mean densities of

sponges on southern California banks were

reported to be 2.5 times higher than corals

(Bright 2007). Sponges on Rittenburg Bank were

particularly abundant, and occurred in larger

sizes and densities than corals (GFNMS 2013).

The most abundant sponges on Piggy Bank

(Yoklavich et al. 2011) and other southern

California banks (Yoklavich unpubl.) generally

were mound, foliose, upright flat, vase, and

barrel morphotypes.

A unique habitat dominated by hexactinellid

sponges has been discovered in the vicinity of

methane seeps off Grays Harbor, Washington

(Salmi et al. 2011). NMFS bottom trawl surveys

in this area in 1983 and 1985 reported

significant catches of sponges, up to 907 kg in a

single ~ 1.5 km trawl haul. Glass sponges

identified from this survey included

Aphrocallistes vastus, Hyalonema sp., and

rossellid sponges. In 2010, the distribution of

the sponges and associated fishes was

quantified during an AUV survey (Clarke and

Fruh 2012). The seafloor was of low relief and

much of it was covered with sediment. On five

dives the densities of sponges ranged from 10

to 99 sponges per 100 m2 and included the

structure‐forming glass sponges Heterochone

calyx, Aphrocallistes sp., and a demosponge,

Poecillastra sp. Many of the rockfishes in the

area were associated with sponges, and a high

percentage of the fishes either were touching or

within one body length of these structure‐

forming sponges.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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Sponge reefs have been described in several

areas of British Columbia (Conway et al. 2001).

Similar to the area off Grays Harbor,

Heterochone calyx and a species of Aphrocallistes

are among the dominant species. However, the

densities of sponges in the Grays Harbor area

are much lower than found at the areas

described as sponge reefs in British Columbia,

Canada. In those areas, densities can typically

be on the order of 2000 per 100 m2 (Leys et al.

2004) while off Grays Harborthe densities

ranged up to 99 sponges per 100 m2.

II.3. Species Associations with

Deep-Sea Corals and Sponges

While demersal fishes co‐occur with various

deep‐sea corals and sponges, particularly in

rocky areas, the extent and function of such

relationships have yet to be fully described and

quantified. Fishes occasionally have been seen

resting inside, underneath, and beside corals

and sponges during visual surveys conducted

with manned submersibles, ROVs, and AUVs

off the west coast (Brancato et al. 2007, Bianchi

2011, Clarke and Fruh 2012). Such associations,

particularly of rockfishes (Sebastes spp.) with

some of the same species of corals or sponges,

have also been reported off British Columbia

(Du Preez and Tunnicliffe 2011) and Alaska

(Stone and Shotwell 2007, Stone 2014, Rooper et

al. 2007). However, there was little evidence of

a functional relationship between fishes and

structure‐forming invertebrates that co‐

occurred in various habitats throughout

southern California (Tissot et al. 2006). Bright

(2007) documented the co‐occurrence of several

species of fishes with structure‐forming

invertebrates on the “Footprint” of the Channel

Islands off southern California. In recent visual

surveys conducted using an ROV and manned

submersible on nearby seamounts in southern

California, there were few instances of fishes

associated with corals and sponges, and benthic

invertebrates more commonly occurred on

dead Antipathes dendrochristos (Christmas tree

black corals) and Heterochone calyx (goblet

sponges) than on living ones (Yoklavich et al.

2011, Yoklavich et al. 2013). Bianchi (2011) also

reported few fishes in close proximity with

structure‐forming invertebrates (including

corals and sponges) in submarine canyons off

Washington and central California, with most

fishes occurring ≥1 m away from the

invertebrate. The extent of fish and invertebrate

associations may depend on the densities of

fishes and structure‐forming invertebrates and

on the local availability of rocky crevices in

which fishes take shelter. Further research and

critical evaluation are needed to clarify the role

that corals and sponges serve as essential

habitat for fishes in demersal communities off

the West Coast.

II.4. Mapping Potential Distribution

of Deep-Sea Corals and Sponges

One of the impediments to identifying areas

that are likely to have abundant populations of

corals and sponges is a lack of high‐resolution

bathymetry, backscatter, and associated

interpreted substratum types on the seafloor in

deep water on a coast‐wide scale. A significant

amount of new information on physical habitat

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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has been collected off the Pacific Coast over the

last decade, including at least 261 sources of

seafloor imagery complied since the last

regional habitat maps were published in 2005

(NMFS 2013). Some of this information has

been used by NOAA to target coral and sponge

surveys. Most of the new information has been

consolidated into Pacific coast‐wide

comparative maps of bathymetry, acoustic

coverage, seafloor substrata, and biogenic

habitat observations in 2005 and 2011 for the

EEZ off Washington, Oregon, and California in

support of the PFMC 5‐year review of EFH for

Pacific coast groundfish (PFMC 2012).

The Office of Ocean Exploration mapped the

Mendocino Ridge and specific areas in and

around the Gulf of the Farallones, Cordell Bank,

and Monterey Bay national marine sanctuaries.

A new ridge southwest of Santa Rosa Island in

the Channel Islands National Marine Sanctuary

was identified during these multibeam surveys

conducted from the NOAA R/V Okeanos

Explorer. NOAA explored this newly charted

ridge and to evaluate the depth stratification of

deep‐sea corals. Beginning in 2015, NOAA

began a series of cruises designed to

comprehensively map the deeper waters of the

Channel Islands National Marine Sanctuary.

NOAA has mapped additional areas in

preparation for targeted ROV or AUV surveys.

Additional bathymetric mapping in the

Olympic Coast and Gulf of the Farallones

National Marine Sanctuaries was completed.

Multibeam surveys were conducted in

preparation for AUV surveys of a sponge area

off Grays Harbor, Washington and off Oregon.

In addition to these federally supported efforts,

all of the seafloor within California’s state

waters (to 3 miles offshore) and large portions

of Oregon state waters have been mapped with

high‐resolution multibeam sonar. These data

have been coupled with predictive models to

map the occurrence of deep‐sea corals.

II.5. Research Priorities

Whitmire and Clarke (2007) identified a

number of deep‐sea coral research priorities for

the region. In 2010, NOAA’s Deep Sea Coral

Research and Technology Program held a

workshop (NOAA 2010) that supported and

expanded on these research priorities for the

West Coast (see below). While research since

2007 has made progress in many of these areas,

most of the identified priorities remain.

1) Identify deep‐sea coral and sponge species, and

map and characterize their distribution,

abundance, densities, and diversity throughout

the California Current LME.

 A great deal of new information is now

available on coral presence, but very

few studies report abundances and

densities in ways that can be compared

across studies. Information on the

identification of sponges at the species

level is particularly weak.

 The lack of coast‐wide seabed mapping

information at a spatial scale relevant to

demersal communities (i.e., meter

resolution) remains an impediment for

predictive models of habitat suitability

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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and of coral and sponge distribution

and abundance. Improvements to

seafloor maps also will yield efficient

design of surveys to target areas with a

high likelihood of containing abundant

populations of corals and sponges.

Additionally, much of the existing

interpreted habitat mapping needs

validation.

2) Determine the ecological roles of deep‐sea corals

and sponges, especially the nature of

associations between other invertebrates and

fishes with deep‐sea corals and sponges.

 While co‐occurrence of sponges and

corals with some species of fishes and

other invertebrates has been

documented, the extent and nature of

those associations has not been

addressed. This is especially important

from a management standpoint in

relation to roles that structure‐forming

species might play as EFH for demersal

fishes.

3) Understand the basic biology of deep‐sea

corals, including taxonomy, age structure,

growth, gender, population connectivity,

and life histories.

 There have been few new studies on the

life history of west coast deep‐sea corals.

Flint et al. (2007) reported on the

reproductive ecology of a deepwater

solitary scleractinian coral, and Feehan

and Waller (2015) examined

reproduction of seven gorgonian species

(P. pacifica, Calcigorgia beringi and five

Swiftia spp.). Andrews et al. (2009) and

Hill et al. (2011) reported on growth and

age of bamboo corals (Family: Isididae)

from seamounts off California.

Estimates of bamboo coral ages ranged

from 98 ± 9 years (Andrews et al. 2009)

to over 300 years (Hill et al. 2011).

 As in most other areas, information on

the biology of deep‐sea sponges lags far

behind that of deep‐sea corals.

4) Develop models to predict distribution and

abundance of deep‐sea corals and sponges.

 There is a particular need to incorporate

bottom‐substratum information into

region‐wide models and to model corals

at the species level (rather than broad

groupings of species).

 To date there have been no models of

deep‐sea sponge distributions off the

west coast.

 Better understanding of abiotic and

biotic habitat requirements will not only

improve habitat‐suitability modeling,

but will result in models that better

predict impacts of changing ocean

conditions due to acidification and

climate change.

5) Understand anthropogenic and natural impacts

on deep‐sea coral ecosystems.

 Develop baseline indices of health and

condition of corals and sponges at

individual and community levels.

 Better quantify bycatch of corals and

sponges in commercial fisheries.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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6) Further synthesize and understand existing

information on deep‐sea corals and sponges.

 Bottom trawl surveys and fisheries

bycatch provide valuable information

on corals and sponges, but identification

of these taxa is slow and inadequate.

This will improve as progress is made in

developing methods of genetic

identification.

7) Understand deep‐sea corals’ ability to recover

from a variety of stressors.

 The impacts and recovery rates of corals

and sponges from various stressors are

largely theoretical because it is difficult

to conduct controlled experiments with

these fragile deep‐sea organisms. As

methodologies improve, particularly

relevant to climate change and ocean

acidification, we are hopeful that

experimental studies will offer valuable

insight.

8) Utilize deep‐sea corals to discern past climate

conditions.

 There have been gains in determining

paleoclimate information deep‐sea

corals (Prouty et al., this volume), but

few studies on the west coast.

9) Evaluate the efficacy of existing management

measures.

 Recent analysis suggests a majority of

coral occurrences on the continental

shelf and upper slope are outside

existing conservation areas that are

meant to protect seafloor habitats from

interactions with certain bottom‐contact

fishing gears (NMFS 2013). In contrast,

MPAs on the lower slope appear to

protect a large majority of known coral

(excluding pennatulaceans) and sponge

habitats from adverse impacts of bottom

trawling. We need to evaluate the

overall efficacy of all management

measures meant to protect deep coral

and sponge ecosystems.

10) Improve tools and methods for studying deep‐sea

corals (e.g., new technologies; new applications

for existing technologies; and develop best

practices for research).

 DNA analysis has become a standard

tool to assist in species identification

and collaborations between observer,

survey and research programs are

helping to increase the number of

samples of corals and sponges available

for analysis (e.g., Everett et al. 2016).

III. Update on Management

of Deep-Sea Corals and

Sponges

III.1. New Information on Impacts

and Stressors

III.1.i – Fishing

Fishing activities were highlighted in the 2007

report as one of the most significant potential

stressors to deep‐sea coral and sponge

communities, and impacts from bottom‐contact

gears, particularly bottom trawls and longlines,

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

remain the most significant documented threat.

Recent and ongoing activities by management

agencies have focused on minimizing these

impacts; however, quantifying these impacts is

difficult. Bycatch (e.g., NMFS 2016) and in situ

evidence of fishing gears entangling corals and

sponges (e.g., Brancato et al. 2007, Bowlby et al.

2011a, 2011b) continue to be the most direct

evidence of adverse fishing interactions. As

observer coverage increases and as fishery

observers identify corals and sponges more

consistently, bycatch data could be a reliable

indicator of trends in fishing impacts.

There have been few in situ studies off the West

Coast that relate observations of trawling to

deep‐sea coral or sponge habitats. Graiff (2008)

analyzed the megafaunal invertebrate

communities at three sites on the continental

shelf and upper slope off central California, and

examined the potential effects of fishing. It was

difficult to discern fishing impacts from

environmental effects on distribution of

invertebrates in part because of the fairly course

resolution of fishing effort information. Hixon

and Tissot (2007) compared benthic

communities in trawled and untrawled mud

habitat off Oregon. Sea pens (Stylatula spp.)

dominated untrawled bottoms, but were nearly

absent on trawled bottoms. Hannah et al. (2010)

also found that densities of sea pens (Halipteris

spp.) were significantly lower in areas off

Oregon that were heavily‐trawled for shrimp,

compared to lightly‐trawled areas. To more

clearly quantify impacts, controlled in situ

experimentation may be necessary.

Yoklavich et al. (2017) reported on visual

surveys in areas of longtime bottom trawling

off southern Oregon and northern California to

evaluate the incidence of disturbance and

damage to deep‐sea corals and sponges. Overall

frequency of disturbance was relatively low

(2%). Most of the observed damage was to coral

colonies, particularly to bamboo corals (family

Isididae).

Deep‐sea corals and sponges are recorded as

bycatch in both the commercial trawl fishery

and fixed gear fisheries (as documented by the

West Coast Groundfish Observer Program).

Taxonomic identification of commercial bycatch

is usually limited to “sea pens and sea whips”

(i.e., pennatulaceans), “other coral,” and

“sponge.” These observations document

ongoing interactions between commercial

fishing gear and deep‐sea corals and sponges.

Since June 2006, observers of the bottom trawl

fishery have recorded a doubling of encounters

with sponges and pennatulaceans, while the

frequency of occurrence of other corals

remained unchanged (Table 4, PFMC 2012).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

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Table 4. (Modified from PFMC 2012.) Summary of coral and sponge bycatch for observed tows using bottom

trawls as part of the West Coast Groundfish Observer Program, comparing two time periods: “Before” (3 Jan

2002 – 11 Jun 2006) and “After” (12 Jun 2006 – 31 Dec 2010) implementation of Amendment 19 regulations.

“#” denotes number of hauls; “FREQ” denotes ratio of hauls with positive catch of taxon to total hauls

observed; “Weight” denotes catch (kg); “CPUE” denotes catch per unit effort (kg/km). Haul counts represent

only those hauls where corals or sponges were present in the catch. Pennatulaceans are recorded separately

from other corals, as they are relatively easy to distinguish and generally occur in different habitats. Annual

Observer Program coverage of the limited‐entry bottom trawl sector can be found online at:

http://www.nwfsc.noaa.gov/research/divisions/fram/observer/sector_products.cfm.

BEFORE AFTER

TAXON # FREQ Weight CPUE # FREQ Weight CPUE

coral 319 2.0% 4,223 2.2E‐02 335 1.8% 997 4.1E‐03

sea pen 198 1.3% 105 5.5E‐04 474 2.5% 66 2.7E‐04

sponge 469 3.0% 4,547 2.4E‐02 1,444 7.6% 20,585 8.5E‐02

Combined 903 5.7% 8,875 4.7E‐02 2,003 10.5% 21,648 8.9E‐02

Also, sponge bycatch appears to have increased

corals from sponges. Other possible

explanations of these bycatch patterns are

changes in fishing patterns due to spatial

management of the trawl fishery and better

recording of bycatch due to increased interest

in conservation of these organisms. Fixed

bottom‐contact fishing gears, such as bottom‐

set longlines and gill nets can also impact coral

and sponge habitats, though the area contacted

is likely to be much smaller than for bottom

trawls. Surveys in the Olympic Coast National

Marine Sanctuary have documented impacts to

biogenic habitats from bottom longline fishing

gear (Brancato et al. 2007, Bowlby et al. 2011a,

2011b). For fixed gear fisheries, corals and

sponges are recorded in the catch infrequently

by the West Coast Groundfish Observer

Program (PFMC 2012), so it is difficult to

examine trends in bycatch. The transition to full

observer coverage with implementation of

catch shares in the groundfish fishery should

someday afford a more meaningful analysis of

bycatch trends.

For the five‐year review of Pacific Groundfish

EFH conducted by NMFS and the PFMC, data

were compiled on fishing effort in three major

gear sectors: bottom trawl, midwater trawl and

fixed gears (see PFMC 2012). The intensity and

extent of fishing effort for each gear type was

mapped for two time periods bracketing

implementation of EFH regulatory measures in

June 2006. Mapping of bottom trawl intensity

showed discrete areas where fishing effort

either increased or decreased after June 2006,

but in general effort appears to have moved

from the continental shelf to deeper slope

habitats. This shift in effort to deeper waters

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

was most likely caused not by EFH regulations

but by the rockfish conservation area, a time‐

area closure designed to reduce bycatch of

overfished groundfish. In fact, the analysis

showed that EFH conservation areas did not

significantly displace bottom trawling primarily

because little (< 5% per year) bottom trawling

occurred within these areas for the 4.5 years

prior to implementation (NMFS 2013). In

addition to the trawl restrictions in the rockfish

conservation area, a footrope size restriction in

late 2000 likely altered behavior of the bottom

trawl fleet away from high‐relief rocky habitats

(Hannah 2003, Bellman et al. 2005). It is these

types of habitats that many of the EFH closures

were designed to protect.

Midwater, or pelagic, trawls do occasionally

contact the seafloor, but observers have

recorded very little bycatch of corals or sponges

since records began in 2000. In fact, corals and

sponges have been recorded in only 0.4% of

tows between 2000 and 2010, totaling only 38.4

kg of bycatch over that time frame (PFMC

2012).

For fixed gears (e.g., longlines, pots, other

hook‐and‐line), the annual proportion of effort

within areas now closed to bottom trawls has

fluctuated between 4 and 18% (NMFS 2013).

This and the fact that observer coverage of

many fixed gear sectors was relatively low

compared to the trawl sector during this time

period suggests it may be difficult to discern

changes to fishing behavior due to

implementation of EFH closures.

III.1.ii – Other Stressors

In addition to fishing, stressors such as

sedimentation caused by oil and gas

exploration, development and deployment of

gas pipelines and communication cables,

mining, pollution, and climate change were

identified in the 2007 report. These continue to

be important when considering the health and

sustainability of deep‐sea corals and sponges;

however, little new information exists on

specific impacts. A moratorium on new leases

for offshore drilling for petroleum has existed

in both federal (since 1984) and California state

waters (since 1969); however, drilling continues

on existing leases. A number of new submarine

telecommunication cables have been installed

in the region since 2007; however, the single

study (Brancato and Bowlby 2005) and other

anecdotal evidence suggest little impact on

deep‐sea corals and sponges. Finally, wind and

wave energy projects have been considered

along the West Coast, with a pilot wave energy

facility permitted off Reedsport, Oregon. This

facility underwent an extensive review by both

state and federal agencies to ensure little impact

to sensitive habitats. In the end, the installation

company abandoned the project for lack of

funding. Other sites would also be nearshore

and should warrant a similar review, as

deployment and operational aspects of these

projects can have significant impact on the

seabed, and should be evaluated for their

potential impact on deep‐sea corals and

sponges (Boehlert et al. 2013). Despite growing

interest from the energy sector, a moratorium

on new federal wave energy permits for

Oregon was put in place in 2008.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Since 2007, the specific role of climate change in

ocean acidification has become more evident.

Ocean acidification is predicted to affect corals

worldwide, but maybe more immediately in the

North Pacific (Guinotte et al. 2006). In the

California Current System, upwelling brings

acidified water up along the continental shelf,

reaching the surface at some locations (Feely et

al. 2008). Aragonite saturation levels (ΩA) on

the shelf may be among the lowest in the world

(Manzello 2010), which is expected to impact

calcification rates for many corals. There have,

however, been no direct studies on the effects of

acidification on west coast corals. Teck et al.

(2010) surveyed 107 experts to quantitatively

estimate the relative vulnerability of California

Current ecosystems. For hard slope ecosystems,

those most likely to support deep‐sea corals

and sponges, the stressors with the highest

scores were ocean acidification and “demersal

destructive fishing.” A recent threat assessment

for the Davidson Seamount Management Zone

identified ocean acidification, sea temperature

rise, and vessel traffic (especially oil tankers) as

the highest level threats to the seamount’s

resources (NOAA 2012).

III.2. New or Planned

Management Actions

Since 2007, a few management measures have

been enacted to expand protection of areas

where corals and sponges likely occur. The

most significant action occurred in 2009, when

the Monterey Bay National Marine Sanctuary

was expanded to include Davidson Seamount

off the coast of central California (15 CFR §

922.130). This undersea mountain, whose crest

is approximately 1,250 m below the sea surface,

is home to a diverse assemblage of dense,

extensive deep‐sea corals and sponges (Clague

et al. 2010). The seamount was previously

protected (in 2006) from fishing with bottom

contact gear (or any other gear) below 3,000 feet

(~ 1,000 m) by NMFS and the PFMC. Sanctuary

designation provided additional protection

against activities that might impact biological

or non‐biological sanctuary resources below

3,000 feet within the 2,006 km2 Davidson

Seamount Management Zone is prohibited

without a permit.

In 2015, NOAA expanded the boundaries of

Cordell Bank National Marine Sanctuary and

Gulf of the Farallones National Marine

Sanctuary (renamed Greater Farallones) to an

area north and west of their old boundaries.

This expansion eliminated the threat from

petroleum development among other potential

impacts to an area that includes major

underwater features, such as Bodega Canyon,

which could harbor significant coral and

sponge resources.

The State of California, under the Marine Life

Protection Act, has designated 94 MPAs in state

waters with varying levels of protection, all of

which comprise a network of protection along

the entire California coast. These areas are

enhancing the protection of corals and sponges

in California State waters, and associated

monitoring and research in these areas has

added to our knowledge of the distribution of

corals and sponges (Starr and Yoklavich 2008,

Yoklavich et al. 2010).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

As part of the PFMC 5‐year review of Pacific

coast groundfish EFH, a large amount of new

data has been summarized and has greatly

expanded our knowledge of the distribution of

corals and sponges in the region (PFMC 2012,

NMFS 2013). This review includes maps of the

boundaries of current MPAs off the west coast.

As part of this review, the Council received

proposals from various sectors of the public to

modify Pacific groundfish EFH. Most of these

proposals include goals to protect deep‐sea

coral and sponge habitats.

The Magnuson‐Stevens Fishery Conservation

and Management Reauthorization Act of 2006

(P.L. 109‐479) directed NOAA to implement a

Deep‐sea Coral Research and Technology

Program and allowed the designation of deep

coral zones. It also gives regional fishery

management councils new discretionary

authority to protect corals and minimize

interactions with fishing gears. This program

funded research that has added to the

information on corals and sponges on the West

Coast. That information combined with a

variety of continuing research projects

conducted by other institutions has enhanced

our knowledge of these resources. In addition,

NMFS recently has developed a Habitat

Assessment Improvement Plan (HAIP) (NMFS

2010) that outlines research priorities and

procedures to improve habitat assessments.

Many research activities associated with the

HAIP will provide information to aid in our

understanding and protection of deep‐sea

corals and sponges.

IV. Conclusions

Since 2007 there has been significant progress in

gathering and organizing information on

seafloor habitats and species, which has

advanced our understanding of deep‐sea coral

abundance and distribution off the U.S. west

coast. Most recently, much of this has been

catalyzed by NOAA’s Deep Sea Coral Research

and Technology Program and data efforts

associated with the PFMC 5‐year review of

Pacific coast groundfish EFH. Deep‐sea corals

on hard substrata are widely distributed and

the composition of assemblages changes with

depth. The dominant species in many areas are

small and occur at relatively low densities, but

in certain areas (e.g., ridges of seamount and

certain banks, particularly in the Southern

California Bight), larger gorgonian species may

occur in patches of high density. Since 2007,

there has been increased interest in sea pens,

which occur on soft sediments and therefore are

much more widespread than other corals.

Information on deep‐sea sponges is much more

rudimentary than our understanding of corals,

and the data on sponges in this chapter is

cursory at best. Most sponges, when recorded

at all, are often not identified below the Phylum

level. As elsewhere in the world, deepwater

species are primarily glass sponges (Class

Hexactinellida) and demosponges (Class

Demospongiae). Still, it appears that in many

hard‐bottom habitats in deeper waters, sponges

are more abundant than corals. Some habitats

are dominated by sponges, such as glass

sponge grounds, while corals are generally

absent. The abundance of sponge habitats is

reflected in their predominant occurrence in

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

catches from the NMFS West Coast Groundfish

Bottom Trawl Survey and in bycatch in bottom

trawl fisheries as recorded by the West Coast

Groundfish Observer Program.

Fishing with bottom‐contact gear, especially

bottom trawls, remains the most immediate

threat to both deep‐sea coral and sponge

ecosystems. While many areas expected to be

prime habitat for corals and sponges were

protected from trawling either indirectly in

2001 or directly in 2006, continued bycatch in

commercial trawl fisheries indicates that

significant interactions remain, and in many

cases are restricted to relatively discrete areas

(Figure 3). Among other stressors, ocean

acidification will be a major long‐term concern

for some corals, although studies of specific

impacts to corals in the region are lacking.

Since 2007, there have been very significant

conservation actions in state waters (e.g., new

MPAs) where corals and sponges are present.

The expansion of Monterey Bay National

Marine Sanctuary to include Davidson

Seamount, and the expansion to create the

Greater Farallones National Marine Sanctuary,

represent the major management actions in the

EEZ since 2007 that serve to protect deep‐sea

coral and sponge ecosystems.

V. Acknowledgements

We appreciate the thoughtful comments on

earlier drafts of this chapter by our three

colleagues: Tom Laidig (SWFSC), Aimee Keller

(NWFSC) and Dan Howard (CBNMS).

Online Annex – Comprehensive list of deep‐

sea corals in the U.S. West Coast region linked

here:

https://deepseacoraldata.noaa.gov/

library/2017‐state‐of‐deep‐sea‐corals‐report

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  

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Appendix 

Table A. Sites of major research expeditions (2007‐2014) referenced in the chapter that included deep‐sea

coral or sponge research. (DSCRTP = Deep Sea Coral Research and Technology Program; MBARI =

Monterey Bay Aquarium Research Institute; NCCOS = National Centers for Coastal Ocean Research; NMS =

National Marine Sanctuary; SWFSC = Southwest Fisheries Science Center; DFO Canada = Department of

Fisheries and Oceans, Canada; BOEM = Bureau of Ocean Energy Management; COPC = California Ocean

Protection Council; CDFW = California Department of Fish and Wildlife; USGS = United States Geological

Survey; NWFSC = Northwest Fisheries Science Center; UW = University of Washington).

Location Year Vessel Description Program

2007

2008

NOAA Ship

David Starr

ROV photo surveys for ESA‐listed abalone

(subsequently analyzed for deep‐sea coral

NOAA SWFSC

Jordan occurrence) on several rocky banks. PI: J. Butler

2008 R/V Velero IV Submersible video surveys for fishes and BOEM

Stylaster californicus on Farnsworth Bank near

Catalina Island. PI: M. Love

Southern

California

Bight &

Channel

Islands NMS

2010

2010

2010

NOAA Ship

McArthur II

R/V Velero IV

CPFV Outer

ROV and AUV surveys of Piggy Bank, Channel

Islands NMS. PI: M. Yoklavich & M.E. Clarke

Submersible surveys of Christmas tree black

coral habitats in Channel Islands NMS. PI: M.

Yoklavich

ROV photo surveys of several rocky banks. PI: J.

NOAA DSCRTP

NOAA DSCRTP

NOAA SWFSC,

2011 Limits Butler & P. Etnoyer NCCOS

2011 R/V Velero IV Submersible surveys of fishes and NOAA SWFSC

invertebrates on rocky banks in Channel Island

NMS. PI: M. Yoklavich

2013 R/V ROV Surveys for deep‐sea coral habitats in NOAA DSCRTP

Shearwater Channel Islands NMS. PI: S. Katz

Pioneer &

2007 R/V Western

Flyer

ROV surveys of Pioneer and Davidson

Seamounts. PI: D. Clague

MBARI

Davidson

Seamounts 2009 R/V Western ROV surveys of Davidson Seamount. PI: J. Barry MBARI

Flyer

2008 R/V Point

Lobos

ROV surveys of Monterey Canyon. PI: C.

McClain

MBARI

2007 R/V Velero IV Submersible video surveys for fishes and COPC, CDFW

2008 invertebrates in and out of 8 offshore MPAs.

Monterey

Bay NMS 2010 NOAA Ship Bell

M. Shimada

PI: M. Yoklavich & R. Starr

ROV photo surveys of Sur Canyon. PI: J. Butler

& P. Etnoyer

NOAA SWFSC,

NCCOS

2010 R/V Derek M.

Baylis

ROV surveys at 17 sites in Monterey and

Carmel Bays. PI: G. Shester

Oceana

2013 R/V Western

Flyer

ROV survey of Sur Ridge. PI: J. Barry & A.

DeVogelaere

MBARI, NOAA

MBNMS

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Table A. continued

Location Year Vessel Description Program

2009 NOAA Ship Multibeam mapping survey of Gulf of the NOAA Ocean

Okeanos Explorer Farallones NMS and potential sanctuary Exploration

Gulf of the

Farallones

NMS

2011 R/V Fulmar

expansion area. PI: M. Malik

Multibeam survey of potential deep‐sea coral

habitats on Rittenburg Bank and Farallon

Escarpment. PI: J. Roletto

NOAA DSCRTP

2012 R/V Fulmar ROV survey of Rittenburg Bank, Cochrane Bank NOAA DSCRTP

and Farallon Escarpment. PI: J. Roletto

2007 R/V Fulmar Camera sled video surveys of deepwater NOAA CBNMS

habitats. PI: D. Roberts

2009 NOAA Ship Multibeam mapping survey of Cordell Bank NOAA Ocean

Cordell Bank

NMS 2010

Okeanos Explorer

NOAA Ship

NMS and potential sanctuary expansion area.

PI: M. Malik

ROV surveys. PI: D. Howard

Exploration

NOAA DSCRTP

McArthur II

2011 R/V Fulmar AUV survey for potential deep‐sea coral NOAA DSCRTP

habitats. PIs: M.E. Clarke & D. Howard

2007‐ R/V Lakota & Camera sled video surveys of seafloor USGS, SWFSC

2012 Coral Sea; R/V invertebrate communities in continental shelf

California

Coast 2014

Fulmar &

Shearwater

R/V Point Sur

habitats throughout California state waters. PI:

G. Cochrane & L. Krigsman

AUV and towed camera surveys of areas of NOAA DSCRTP

high bycatch of deep‐sea corals off Northern

California

2010 NOAA Ship Bell ROV photo surveys of Coquille Bank. PI: J. NOAA SWFSC,

M. Shimada Butler & P. Etnoyer NCCOS

2011 R/V Miss Linda ROV surveys at Cape Arago, Coquille Reef and Oceana

Oregon Coast Bank, Orford Reef. PI: B. Enticknap

2013 R/V Miss Linda ROV surveys at Stonewall Bank, Hecata Bank, Oceana

Daisy Bank, Siletz Hotspot, Siletz Reef/Cascade

Head. PI: B. Enticknap

2007 R/V Thomas G. Side‐scan sonar and Van Veen grab surveys of UW

Grays

Canyon 2009

Thompson

R/V Kvichak

Defender VI

glass sponge grounds. PI: P. Johnson

ROV survey of glass sponges and associated

methane seeps and krill swarms. PI: P. Johnson

UW

2010 R/V Pacific Storm AUV surveys of glass sponges. PI: M.E. Clarke NOAA DSCRTP

2008 CCGS John P.Tully ROV surveys. PI: E. Bowlby NOAA OCNMS,

DFO Canada

2008 NOAA Ship Multibeam mapping survey. PI: M. Malik NOAA Ocean

Okeanos Explorer Exploration

Olympic 2010 NOAA Ship ROV and AUV surveys. PI: E. Bowlby NOAA DSCRTP

Coast NMS McArthur II

2010 NOAA Ship Bell ROV photo surveys. PI: J. Butler & P. Etnoyer NOAA SWFSC,

M. Shimada NCCOS

2011 R/V Pacific Storm Multibeam survey of potential deep‐sea coral NOAA DSCRTP

habitats. PI: E. Bowlby

Cobb

Seamount

2012 CCGS John P.Tully ROV and AUV surveys of corals and sponges.

PI: M.E. Clarke & J. Curtis

NOAA DSCRTP,

NOAA NWFSC,

DFO Canada

Bowie

Seamount

2011 CCGS John P.Tully ROV and AUV surveys for corals and sponges.

PI: M.E. Clarke & J. Boutillier

NOAA DSCRTP,

NOAA NWFSC,

DFO Canada

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Map 1. Locations of black corals (Order Antipatharia) recorded in the National Deep‐Sea Coral and Sponge

Database (as of October 2017). PFMC = Pacific Fishery Management Council.

147
                       

     

                               

          

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Map 2. Locations of stony corals (Order Scleractinia) recorded in the National Deep‐Sea Coral and Sponge

Database (as of October 2017).

148
                       

     

                           

                          

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Map 3. Locations of non‐gorgonian alcyonacean (true soft corals and stoloniferan corals; Order Alcyonacea)

recorded in the National Deep‐Sea Coral and Sponge Database (as of October 2017).

149
                       

     

                             

         

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Map 4. Locations of gorgonians (formerly Order Gorgonacea) recorded in the National Deep‐Sea Coral and

Sponge Database (as of October 2017).

150
                       

     

                               

         

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Map 5. Locations of sea pens (Order Pennatulacea) recorded in the National Deep‐Sea Coral and Sponge

Database (as of October 2017).

151
                       

     

                             

               

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Map 6. Locations of lace corals (Order Anthoathecata, Family Stylasteridae) recorded in the National Deep‐Sea

Coral and Sponge Database (as of October 2017).

152
                       

     

                             

       

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Map 7. Locations of demosponges (Class Demospongiae) recorded in the National Deep‐Sea Coral and Sponge

Database (as of October 2017).

153
                       

     

                               

         

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Map 8. Locations of glass sponges (Class Hexactinellida) recorded in the National Deep‐Sea Coral and Sponge

Database (as of October 2017).

154
                       

     

                           

               

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. WEST COAST

  

Map 9. Locations of sponges (Classes Homoscleromorpha, Calcarea, and unspecified) recorded in the National

Deep‐Sea Coral and Sponge Database (as of October 2017).

155
               

             

           

Black coral Bathypathes on a rocky ridge crest

of Johnston Atoll. Courtesy of the NOAA

Office of Ocean Exploration and Research.
RECENT RESEARCH & MANAGEMENT OF HAWAIIAN BLACK CORALS

• • •

AN UPDATE ON 

RECENT RESEARCH 

AND MANAGEMENT 

OF HAWAIIAN 

BLACK CORALS

I. Introduction

Antipatharians, commonly known as black corals, are a 

little studied order of anthozoan hexacorals that currently 

encompasses over 235 described species (Cairns 2007, Daly 

et al. 2007, Bo 2008). Black corals occur worldwide in all 

oceans from polar to tropical regions, and have a wide 

depth distribution ranging from 2-8,600 m (reviewed by 

Wagner et al. 2012a). Despite this wide bathymetric range, 

black corals are primarily found in deeper waters below 

the photic zone, with over 75% of known species occurring 

exclusively below 50 m (Cairns 2007). At these depths, 

antipatharians are often abundant and dominant faunal 

components, and create habitat for a myriad of associated 

organisms (reviewed by Wagner et al. 2012a).

In Hawai‘i black corals are particularly important, not just 

from an ecological perspective, but also from an economic 

and cultural one. First, black corals are the predominant 

habitat-forming species on Hawaiian deep reefs that are 

exposed to strong currents (50-150 m; Grigg 1965, Grigg 

1976, Chave and Malahoff 1998, Parrish and Baco 2007). 

Second, Hawai‘i is the only place in the United States, and 

only one of a few places in the world, where black coral 

is harvested commercially for the precious coral jewelry 

industry, a multi-million dollar business that employs 

close to 650 people in manufacturing facilities and retail 

stores statewide (Grigg 2001, Parrish and Baco 2007, Grigg 

2010). Third, black corals are important culturally as they 

Daniel Wagner1*, 

Dennis M. 

Opresko2, Anthony 

D. Montgomery3,

and Frank A.

Parrish4

• • •

1 NOAA Papahānaumokuākea 

Marine National Monument

Honolulu, HI

(current affiliation: JHT, Inc., 

NOAA National Centers of 

Coastal Ocean Science

Charleston, SC)

* Corresponding Author:

Daniel.Wagner@noaa.gov

2 National Museum of 

Natural History, Smithsonian 

Institution, Washington DC

3 U.S. Fish and Wildlife Service, 

Pacific Islands Fish and 

Wildlife Office, Honolulu, HI 

4 NOAA Ecosystem Sciences 

Division, Pacific Islands 

Fisheries Science Center, 

Honolulu, HI

157
RECENT RESEARCH & MANAGEMENT OF HAWAIIAN BLACK CORALS

• • •

158

represent the official gemstone of the State 

of Hawai‘i, and were traditionally used in 

Hawaiian culture for medicinal purposes 

(Kaaiakamanu and Akina 1922, Chun 1994). 

‘Ēkaha kū moana, as black coral is known in 

the Hawaiian language, has for a long time 

been cherished by native Hawaiians for its 

presumed therapeutic effects against lung 

diseases and mouth sores (Kaaiakamanu and 

Akina 1922, Chun 1994). 

Despite the fact that native Hawaiians have 

used black corals for several centuries, 

Hawaiian antipatharians were only discovered 

by western scientists at the beginning of the 

20th century. In 1902, the scientific expedition 

by the U.S. Fish Commission Steamer Albatross

was the first to sample black corals from the 

Hawaiian Archipelago (Bayer 1961). Through 

the use of deep-sea trawls and tangle nets, the 

Albatross sampled black corals in Hawaiian 

waters at depths exceeding 2,000 m, although 

the majority of antipatharian specimens 

were collected at depths shallower than 

500 m (Bayer 1961). Black coral specimens 

collected by the Albatross were deposited 

in the National Museum of Natural History, 

where they remained largely unexamined 

until recent reexaminations of these collections 

led to the description of numerous new 

antipatharian species including several from 

Hawaiian waters (Opresko 2002, 2003c, 2005, 

Wagner and Opresko 2015, Wagner 2015a). 

In 1928, Verrill was the first to formally 

describe Hawaiian black coral species, when 

he described Antipathes grandis and Antipathes 

irregularis (Verrill 1928), the latter of which 

was later identified as a gorgonian (Grigg and 

Opresko 1977). In 1958, large aggregations of 

black corals were discovered off Maui (Gage 

1962, Grigg 1964), a discovery that eventually 

led to the development of a local black coral 

fishery (Grigg 2001), as well as to numerous 

studies on this group of corals. Scientists, 

resource managers and fishermen have 

collaborated closely throughout the history 

of the fishery (Grigg 2001), and as a result, 

Hawaiian antipatharians are among the best 

studied on the globe (Etnoyer and Morgan 

2005, Wagner et al. 2012a). The purpose of 

this article is to briefly summarize these 

collaborations, with an emphasis on black 

coral research and management activities 

that have occurred since the last synthesis on 

the state of the deep-sea coral ecosystems in 

Hawai‘i (Parrish and Baco 2007).

Figure 1. (a) Map showing the geographic locations of the three largest commercially valuable black coral 

beds in Hawai‘i. (b) Diver attaching a black coral colony to an anchor line so it can be lifted to the surface 

(Photo by R. Grigg).
RECENT RESEARCH & MANAGEMENT OF HAWAIIAN BLACK CORALS

• • •

159

II. The History of the Hawaiian

Black Coral Fishery

Since its inception in 1958, the Hawaiian black 

coral fishery has been supplied by SCUBA 

divers that collect the coral at depths between 

40-70 m, primarily in the Au‘au Channel

between the islands of Maui and Lāna‘i, and

to a lesser extent off the islands of Kaua‘i and

Hawai‘i (Figure 1; Oishi 1990, Grigg 2001,

Parrish and Baco 2007). Harvesting methods

have remained essentially unchanged since

start of the fishery, and involve fishermen

descending to black coral beds using SCUBA,

severing colonies from the substrate using

axes or sledges, and then bringing coral trees

back to the surface via lift bags or anchor

lines (Figure 1; Oishi 1990, Grigg 2001). Three

black coral species have been reported as

being targeted by the Hawaiian precious coral

fishery, and include Antipathes griggi Opresko,

2009 (formerly A. dichotoma), Antipathes grandis

Verrill, 1928, and Myriopathes cf. ulex (Ellis

and Solander, 1786) (formerly Antipathes ulex)

(Figure 2; Grigg 1976, NOAA 1983, Oishi

1990, DLNR 1999, Grigg 2001, Parrish and

Baco 2007, Grigg 2010). However, close to

90% of the coral harvested in Hawaiian waters

consists of A. griggi, with A. grandis and M.

cf. ulex representing 10% and 1% of harvested

black coral, respectively (Oishi 1990).

III. Black Coral Fishery

Management

The vast majority of commercially valuable 

black coral beds lie in waters that fall under 

the jurisdiction of the State of Hawai‘i (< 3 

nautical miles from shoreline), with a much 

lesser degree found in federal waters (3-200 

nautical miles from shore) (Figure 1; Grigg 

2001, Parrish and Baco 2007, Grigg 2010). 

The fishery is therefore managed through 

both state and federal regulations, which are 

set by the Hawai‘i Department of Land and 

Natural Resources (DLNR), and NOAA’s 

National Marine Fisheries Service (NMFS) in 

consultation with the Western Pacific Regional 

Fishery Management Council (WPRFMC), 

respectively (NOAA 1983, DLNR 1999, 

NOAA 2002, DeMello 2006, NOAA 2007, 

2008). State regulations came into effect in 

1981 (DLNR 1999). Since then, all harvesters 

of black corals in Hawaiian state waters are 

required to purchase a state commercial 

marine license, and submit catch reports if 

the coral is sold (Oishi 1990). In 1999, the 

State of Hawai‘i added size limits by which it 

prohibited harvesting of colonies with a base 

diameter of less than 1.905 cm (DLNR 1999), 

which corresponds to a colony height of 90 

cm (Boland and Parrish 2005), or an age of 

14-15 years using growth rate estimates (6.12-

6.42 cm/yr; Grigg 1976). Federal regulations

on the fishery became effective in 1983,

which mandated domestic fishermen to

report their catches in federal waters (NOAA

1983). In 2002, federal regulations added size

harvesting limits, which prohibited harvesting

of colonies with a base diameter of less than

2.54 cm or a colony height lesser than 122 cm

(NOAA 2002). Additionally, the 2002 federal

rule provided persons that had reported black

coral landings between 1997 and 2002 the

option to apply for an exemption, that allowed

harvesting using the minimum base diameter

of 1.905 cm (NOAA 2002). This exemption

was removed in 2007, thereby effectively

prohibiting all harvesting of colonies less

than 122 cm in height or 2.54 cm in width

from federal waters (NOAA 2007). In 2008,

a harvesting quota of 5,000 kg/2 years was

established for the Au‘au Channel bed (Figure

1), where the vast majority of black corals

have been harvested since 1980 (Parrish 2006,

NOAA 2008). In 2012, an annual catch limit of
160

2,500 kg/year was set for the Au‘au Channel 

bed (NOAA 2012). While this annual catch 

limit applies to the entire Au‘au Channel 

bed, which extends through both state and 

federal waters, it is currently only enforced 

in federal waters (NOAA 2012). Additionally, 

the 2012 revisions set an annual catch limit 

of 1,000 kg/year for all precious corals in the 

Hawai‘i exploratory area (NOAA 2012). This 

catch limit applies to all precious corals found 

in federal waters anywhere else outside the 

Au‘au Channel bed in Hawai‘i, and includes 

pink, red, bamboo, gold (currently under 

moratorium) and black corals (NOAA 2012).

IV. Recent Research

The recent revisions of fishery regulations 

(NOAA 2007, 2008, 2012), were the result 

of a number of deep-water surveys (40-110 

m) that indicated a decline in the biomass of

commercially valuable black corals (Grigg

2001, 2004). Surveys performed in the Au‘au

Channel in 1975 and 1998, showed similar

colony size-frequency distributions, and

thereby suggested a long-term stability

in the recruitment and growth of black

coral populations (Grigg 2001). In contrast,

subsequent surveys performed in 2001,

revealed a downward shift towards smaller

sized colonies, as well as a 25% decline in

the biomass of black corals since 1998 (Grigg

2004). Likely causes for these declining

patterns included both increases in harvesting

pressure, and overgrowth of black corals by

the invasive octocoral Carijoa sp. (initially

identified as Carijoa riisei; Grigg 2003, 2004,

Kahng and Grigg 2005). Together, these

developments raised questions about whether

fishery regulations needed to be redefined

in order to maintain a sustainable harvest

(Grigg 2004). To address these concerns, the

WPRFMC held a workshop to review the state

of the Hawaiian black coral fishery and to 

identify future research priorities (WPRFMC 

2006). The workshop highlighted the need for 

studies on the (1) growth, (2) reproduction, 

(3) recruitment, (4) mortality, (5) fishing, and

(6) geographical distributions of Hawaiian

antipatharians (WPRFMC 2006). Motivated in

large part by these recommendations, several

studies have recently examined various

aspects of the taxonomy, biology and ecology

of commercially valuable Hawaiian black

corals, and are briefly reviewed below.

IV.1. Taxonomy

In 1977, Grigg and Opresko (1977) published a 

taxonomic survey of Hawaiian antipatharians 

based on colony branching pattern. Since that 

study, skeletal spine morphology has become 

increasingly important in antipatharian 

taxonomy (Opresko 2001, 2002, 2003c, 2004, 

2005, 2006, and references therein), because 

this character is thought to be largely 

independent of environmental cues, as 

compared to other more plastic morphological 

characters (Lapian et al. 2007, Wagner et al. 

2010). Recent taxonomic studies using skeletal 

spine morphology have led to more detailed 

descriptions of several Hawaiian species, as 

well as to revisions of the species names that 

had previously been used in the literature 

(Opresko 2009, Wagner et al. 2010, Wagner 

2011, Wagner et al. 2011a, Opresko et al. 2012, 

Wagner 2015a). While these recent revisions 

have provided better insights into what 

species are present in Hawaiian waters and 

how they compare to species found elsewhere, 

systematics of the entire antipatharian order 

are still in a state of revision. 

Prior to recent revisions, the Hawaiian 

species Antipathes griggi (Figure 2) was 

known as A. dichotoma Pallas, 1766, a species 

originally described from off Marseilles in the 

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161

Figure 2. Hawaiian antipatharian species (a) Antipathes griggi, (b) Antipathes grandis, (c) 

Myriopathes cf. ulex, and (d) Aphanipathes verticillata (scale bars = 200 μm). NOTE: With the 

exception of M. cf. ulex, species cannot be reliably differentiated based on colony morphology alone, 

and typically require microscopic examination of skeletal spines (bottom row). Photos courtesy of the 

Hawaiian Undersea Research Laboratory (HURL).
162

Mediterranean Sea (Pallas 1766). Subsequent 

comparisons between Hawaiian and 

Mediterranean specimens revealed substantial 

morphological differences (Opresko 2003a). 

As a result, the Hawaiian “A. dichotoma” was 

assigned the new name of Antipathes griggi

and formally described (Opresko 2009).

Even though Antipathes grandis (Figure 2) was 

the first antipatharian species to be described 

from Hawai‘i (Verrill 1928), the original 

species description was very brief and the 

type material remained unexamined for a 

long period. Therefore, Wagner et al. (2010) 

redescribed A. grandis using morphological 

and molecular characters, in conjunction 

with in situ observations. These examinations 

revealed that A. grandis colonies can exhibit 

three different color morphotypes, at least one 

of which may have been confounded with A. 

griggi in the past (Figure 2). 

Grigg (1964) presented the first published 

account of Myriopathes cf. ulex (Figure 2) from 

Hawai‘i (as Antipathella sp.), and Grigg and 

Opresko (1977) later identified this species 

as Antipathes ulex Ellis and Solander, 1786. In 

2001, Opresko reassigned A. ulex to the newly 

established antipatharian family Myriopathidae

and the new genus Myriopathes, resulting in 

the name Myriopathes ulex. The original species 

description of M. ulex is rather brief (Ellis 

and Solander 1786), and the type material 

has been lost (Opresko 2001). Thus, positive 

identification of M. ulex cannot be made 

until a neotype is designated. Pending such 

a taxonomic revision, the name Myriopathes 

cf. ulex is used here and elsewhere to refer to 

the species that is targeted by the Hawaiian 

black coral fishery (Wagner 2011, Wagner et al. 

2012b, Wagner 2015b). 

Besides these three species that had been 

known to exist in areas where black corals 

have been harvested in Hawai‘i (A. griggi, A.

grandis and M. cf. ulex), recent surveys have

discovered a fourth species, Aphanipathes

verticillata Brook, 1889 (Figure 2), which was

previously unknown from Hawaiian waters

(Opresko et al. 2012). While gross observations

suggest that the skeleton of this species is

suitable to manufacture jewelry and may

therefore have been targeted by the fishery, all

known records of Hawaiian A. verticillata come

from depths that are slightly deeper (88-130

m) than the harvesting depth zone (40-70 m;

Opresko et al. 2012). It is therefore unknown

if, and to what extent, A. verticillata has been

commercially harvested in Hawai‘i (Opresko

et al. 2012).

IV.2. Geographic and Depth

Ranges

Surveys for A. griggi have been particularly 

frequent in Hawai‘i, because it is the main 

fishery species. Based on field observations 

using colony morphology, A. griggi has 

been reported throughout the Hawaiian 

Archipelago to depths of 159 m (Chave and 

Malahoff 1998). However, recent taxonomic 

studies have shown that A. griggi can easily 

be misidentified as the sympatric species A. 

grandis or Aphanipathes verticillata (Figure 2). 

These recent taxonomic studies raise questions 

about the validity of many previous reports 

of A. griggi (as A. dichotoma), especially 

those at depths below 100 m. Additionally, 

colonies with similar morphologies to A. 

griggi, identified as A. dichotoma, have been 

reported from various Indo-Pacific locations 

including the Philippines, Indonesia, Palau, 

China, Guam and Johnston Atoll (Van Pesch 

1914, Grigg 1975, Zhou and Zou 1984, Zou 

and Zhou 1984, Zhou and Zou 1992, Chave 

and Malahoff 1998, Paulay et al. 2003, Rogers 

et al. 2007, Qi et al. 2009). These locations all 

lie outside the range of A. dichotoma, which 

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163

is only known from the Mediterranean and 

East Atlantic (Opresko 2003a, Bo 2008). 

Like the previous misidentification of A. 

dichotoma from Hawai‘i, these misidentified 

A. dichotoma records may also be A. griggi.

However, detailed taxonomic investigations

of specimens from these Indo-West Pacific

locations will have to be undertaken to

confirm this. To date, specimens identified

as A. griggi using all available diagnostic

characters, have only been reported from the

Hawaiian Archipelago from the islands of

Hawai‘i to Laysan at depths ranging between

10-110 m (Opresko 2009, Wagner 2011, Wagner

et al. 2011a, Wagner 2015a,b).

A. grandis was originally described from a

specimen collected off Maui (Verrill 1928), and

subsequently reported throughout the Main

Hawaiian Islands from Hawai‘i to Ni‘ihau at

depths between 27-127 m (see Wagner et al.

2010). Additionally, there are two reports of

this species from off China (Zhou and Zou

1984, Zou and Zhou 1984); however, these

records cannot be confirmed until specimens

from that locality are examined.

Myriopathes ulex was initially described from 

Indonesia (Ellis and Solander 1786), but 

subsequently reported throughout the IndoWest Pacific at depths ranging between 25-364 

m (Blainville 1834, Gray 1857, Brook 1889, 

Figure 3. Depth distribution by species for 250 black coral colonies sampled at depths between 30-130 m 

in the Au‘au Channel.
164

Van Pesch 1914, Grigg and Opresko 1977, 

Chave and Malahoff 1998, Parrish and Baco 

2007, Rogers et al. 2007, Bo 2008, Moon and 

Song 2008). However, a thorough taxonomic 

investigation is needed to verify whether these 

records all correspond to the same species 

that is present in Hawaiian waters. In the 

Hawaiian Islands, Myriopathes cf. ulex has 

been confirmed from Hawai‘i Island to Pearl 

and Hermes Atoll at depths ranging between 

41-326 m (Wagner 2011, Wagner et al. 2011a,

Wagner 2015b).

Up to now the black coral fishery has been 

managed under the presumption that a 

harvesting depth refuge exists; however recent 

studies have shown that there is no such depth 

refuge (Wagner et al. 2012b). Due to logistical 

constraints of SCUBA diving, fishermen have 

traditionally harvested black corals at depths 

between 40-70 m. However, dense black 

coral populations exist in the Main Hawaiian 

islands to depths of at least 110 m (Grigg 2001, 

Parrish and Baco 2007). It had previously been 

thought that colonies below the harvesting 

depth zone (> 70 m) provided a depth refuge 

from the fishery and were capable of reseeding 

fished populations in shallower water (< 70 m; 

Grigg 1976, 2001). However, recent surveys 

indicate that the majority of black corals 

below 70 m consist of A. grandis (68%) and 

Aphanipathes verticillata (25%), with A. griggi

accounting for only 7% (Figure 3; Wagner et 

al. 2012b). Thus, the population size below the 

harvesting depth zone has been overestimated 

in the past and there is no real depth refuge 

from harvest (Wagner et al. 2012b).

IV.3. Sexual Reproduction and

Histological Studies

As many other aspects of the antipatharian 

biology (see Wagner et al. 2012a), information 

on the sexual reproduction is scarce, and is 

mostly limited to brief notes accompanying

taxonomic descriptions (reviewed by Wagner

et al. 2011b). Recent histological examinations

of eight Hawaiian species, including the

commercially valuable A. griggi and A.

grandis, have revealed that there are several

similarities in the sexual reproduction of

black corals, even among distantly related

species (Wagner et al. 2011b, Wagner et al.

2012a). First, of the six primary mesenteries

only the two in the transverse plane bear

gametes. Second, entire colonies are generally

either female or male, although sequential

hermaphroditism cannot be ruled out in most

cases. Third, there is no evidence for internal

fertilization within antipatharians, indicating

that fertilization and larval development likely

occur externally in the water column and not

internally within polyps. Additionally, a more

detailed study on the sexual reproduction of

A. griggi has demonstrated that this species

(1) is gonochoric with a 1:1 sex ratio, (2) has

an annual reproductive cycle, and (3) spawns

in multiple successive events with greatest

intensities between November and December

(Wagner et al. 2012b). Furthermore, this study

revealed that ~80% of colonies meeting the

state harvesting limit (90 cm) are sexually

mature, whereas ~90% of colonies meeting

the federal limit (122 cm) are sexually mature

(Wagner et al. 2012b).

Apart from providing insights into the 

sexual reproduction of antipatharians, 

recent histological investigations have 

uncovered endosymbiotic algae of the genus 

Symbiodinium, also known as zooxanthellae, 

within a large proportion of Hawaiian black 

coral species (Wagner et al. 2011c). Due to the 

predominant occurrence of antipatharians 

in low-light environments that do not 

support photosynthesis (> 50 m; Cairns 

2007), the absence of Symbiodinium within 

black corals has generally been inferred 

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165

rather than empirically demonstrated, 

and the whole taxonomic order has been 

considered azooxanthellate (see Wagner et 

al. 2011c). That said, close to a century ago 

several scientists noted round structures 

within the tissues of several black coral 

species, which they interpreted as symbiotic 

algae (Brook 1889, Van Pesch 1914, Buchner 

1921). However, these early reports have 

been largely overlooked or dismissed as 

questionable. Supporting the tentative nature 

of these reports, numerous more recent 

studies using histological techniques (Grigg 

1964, 1976, Goenaga 1977), chlorophyll 

measurements (Shick and Dykens 1985, 

Santiago-Vazquez et al. 2007) and molecular 

techniques (Santiago-Vazquez et al. 2007) 

have failed to detect Symbiodinium within 

antipatharians. In contrast, Wagner et al. 

(2011c) detected very low densities (0–92 

cells/mm3) of endosymbiotic Symbiodinium

within the majority of black coral species from 

Hawai‘i, and down to a maximum depth of 

396 m. While the physiological role of the 

endosymbiotic cells were not determined, 

their low densities coupled with the extreme 

depths at which they were recorded, argue 

that they do not provide their antipatharian 

hosts with photosynthetic products and thus 

suggest an either parasitic or commensal 

association with black corals (Wagner et al. 

2011c).

IV.4. Overgrowth by Carijoa sp.

The alcyonacean octocoral Carijoa sp. was 

first observed in Hawai‘i in 1966 (Kahng et 

al. 2008) and the first published record of this 

species was in 1972, when it was discovered 

within the fouling community of Pearl Harbor, 

O‘ahu (Evans et al. 1974). In 2001, black 

coral surveys in the Au‘au Channel recorded 

Carijoa sp. overgrowing a large percentage 

of commercially valuable Hawaiian black 

corals at depths between 70-110 m (Figure 4; 

Grigg 2003). Due to the concerns about the 

effects of this invasion on the Hawaiian black 

coral fishery, several studies were initiated 

to characterize and monitor the impacts of 

Carijoa sp. on black corals (Kahng and Grigg 

2005, Kahng et al. 2008, Concepcion et al. 

2010, Kahng 2010). Initial surveys in the 

Figure 4. (a-b) Commercially valuable Hawaiian black corals, overgrown by the invasive octocoral 

Carijoa sp. in the Au‘au Channel. Photos courtesy of HURL.
166

Au‘au Channel indicated that over 75% of the 

number of A. grandis and A. griggi colonies 

were overgrown by the octocoral at depths 

between 80-110 m (Kahng and Grigg 2005). 

However, subsequent monitoring efforts in 

the channel suggested that the impact of the 

Carijoa sp. invasion was not worsening with 

time, and that while still widespread and 

serious, the invasion was less severe than 

previously thought (Kahng 2010). These 

more recent surveys have determined that 

~33% of black corals > 40 cm have Carijoa

sp. overgrowth, with ~25% of colonies being 

completely overgrown and ~8% being partially 

overgrown (Kahng 2010). Furthermore, 

Carijoa sp. appears to settle on portions of 

black coral colonies where the bare skeleton is 

exposed, which can be created by abrasion or 

senescence (Kahng and Grigg 2005). There are 

no known natural predators of Hawaiian black 

corals (Wagner et al. 2012a), and therefore it is 

unknown whether predation facilitates Carijoa 

sp. overgrowth.

V. Future Management

Directions and Implications

for Other Coral Fisheries

While the latest surveys on the effects of 

Carijoa sp. on commercially valuable black 

corals indicate that the impacts are less severe 

than first assumed (Kahng 2010), recent 

studies also indicate that there is no depth 

refuge from harvest (Wagner et al. 2012b). 

Consequently, the biomass of commercially 

valuable black corals will likely continue to 

decline if no corrective management actions 

are undertaken (Grigg 2004). Specifically, 

three revisions in the management regulations 

should be pursued, including (1) increasing 

both the state and federal harvesting size 

limits to 130 cm to ensure that more colonies 

have a chance to reproduce before being

exposed to fishing mortality, (2) setting

aside no-take areas to allow more colonies

to continuously reproduce and reseed fished

populations, and (3) limiting the entry of

new fishermen. While there are only three

permitted vessels in the Hawaiian black coral

fishery (NOAA 2007), advances in diving

technologies may attract more people into

collecting black coral in the near future.

Traditionally, diving for black corals has

been a hazardous occupation due to the

many inherent risks associated with deep

SCUBA diving (Grigg 2001, 2010). However,

as new diving technologies such as mixedgas technical diving and closed-circuit

rebreathers become more readily available,

there will likely be an increased incentive

for people to enter the fishery. Fetching

over $25/lb., harvesting black corals can be a

lucrative enterprise (Grigg 2001). With novel

technologies making black coral harvesting

a safer practice, there will likely be a future

increase in the fishing pressure on black

corals. Thus, limiting the number of people

entering the fishery should be pursued now

before the resource becomes too depleted.

Furthermore, creating no-take areas may

allow for the reseeding of fished populations.

With the exception of black corals found

in federal waters around Hawai‘i, there is

currently no limit on the amount of black

coral that can be harvested in Hawai‘i. As

black coral populations in federal waters get

progressively depleted, fishermen might be

pushed towards exploring black coral beds

closer to shore. Without any other regulations,

these beds are currently vulnerable to

becoming overexploited, especially if new

diving technologies attract more people to

enter the fishery and allow for more efficient

harvesting.

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RECENT RESEARCH & MANAGEMENT OF HAWAIIAN BLACK CORALS

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167

Besides Hawai‘i, black corals have been 

commercially harvested in several other 

regions around the globe including 

throughout Asia, South America, the 

Caribbean, the Mediterranean and the Red 

Sea (Grigg 1975, Noome and Kristensen 1976, 

Castorena and Metaca 1979, Olsen and Wood 

1980, Kenyon 1984, Grigg 1993, Romero 1997, 

Padilla and Lara 2003, Deudin et al. 2010, 

Tsounis et al. 2010). However, in contrast to 

the Hawaiian fishery, antipatharian fisheries 

in most other locations have remained largely 

unmanaged and thus exhibited a cyclic pattern 

of discovery of a population, exploitation, 

depletion, followed by exploration for new 

harvesting grounds, a boom and bust cycle 

that resembles strip mining more than 

a fishery (Noome and Kristensen 1976, 

Castorena and Metaca 1979, Grigg 1993, 

Romero 1997, Padilla and Lara 2003, Tsounis 

et al. 2010). In contrast, the Hawaiian black 

coral fishery has had a comparatively long 

history and serves as a commendable example 

of what can be achieved through close 

collaborations between resource managers, 

scientists and fishermen. Through continued 

research partnerships between different 

fishery stakeholders, black coral populations 

in Hawai‘i have become some of the best 

studied in the world, and it is imperative that 

adaptive management strategies continue 

to be pursued as new scientific information 

becomes available. 

VI. Literature Cited

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investigations to date. Hawaiian Shell News 9:8

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zoophytologie, Paris

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assemblages in the black coral beds off 

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420

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EC, Coles SL, Toonen RJ (2010) Resolving 

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CS, Opresko DM, Rodriguez E, Romano SL, 

Stake JL (2007) The phylum Cnidaria: A review 

of phylogenetic patterns and diversity 300 

years after Linnaeus. Zootaxa 1668:127-182
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science and management workshop report. 

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resources. Chapter 91. Pink, gold, and black 

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many curious and uncommon zoophytes 

collected by the late John Ellis, systematically 

arranged and described by the late 

Daniel Solander, London

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deep-sea corals in the Northeast Pacific Ocean. 

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Jokiel PL, Kam DT, Kay EA, Peeling TJ, Smith 

SV (1974) Pearl Harbor Biologica Survey - Final 

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but with vastly different sequels. Hawaiian 

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(Zoantharia; Antipatharia) with 

observations on aspects of their biology. M.S. 

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Mayaguez, Puerto Rico

Gray JE (1857) Synopsis of the families and genera 

of axiferous zoophytes or barked corals. 

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London 25:278-294

Grigg RW (1964) A contribution to the biology and

ecology of the black coral, Antipathes 

grandis in Hawai‘i. M.S. M.S. thesis, University 

of Hawai‘i, Honolulu, Hawai‘i

Grigg RW (1965) Ecological studies of black coral 

in Hawai‘i. Pacific Science 19:244-260

Grigg RW (1975) The commercial potential of 

precious corals in the western Caroline Islands, 

Micronesia. Sea Grant Technical Report. 

UNIHI-SEAGRANT-AR-75-03

Grigg RW (1976) Fishery management of 

precious and stony corals in Hawai‘i. Sea 

Grant Technical Report. UNIHI-SEAGRANTTR-77-03

Grigg RW (1993) Precious coral fisheries of Hawai‘i 

and the U.S. Pacific Islands. Marine Fisheries 

Review 55:50-60

Grigg RW (2001) Black coral: history of a 

sustainable fishery in Hawai‘i. Pacific Science 

55:291-299

Grigg RW (2003) Invasion of a deep coral bed by an 

alien species, Carijoa riisei, off Maui, 

Hawai‘i. Coral Reefs 22:121-122

Grigg RW (2004) Harvesting impacts and invasion 

by an alien species decrease estimates on black 

coral yield off Maui, Hawai‘i. Pacific Science 

1:1-6

Grigg RW (2010) The precious corals fishery 

management plan of the Western Pacific 

Regional Fishery Management Council. Pacific 

Islands Fishery Monographs 1:1-9

Grigg RW, Opresko DM (1977) Order Antipatharia, 

black corals. In: Devaney DM, Eldredge LG 

(eds) Reef and Shore Fauna of Hawai‘i Section 

1: Protozoa through Ctenophora. Bishop 

Museum Press, Honolulu

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of medicinal value found among the 

mountains and elsewhere in the Hawaiian 

Islands, and known to the Hawaiians 

to possess curative and palliative properties 

most effective in removing physical 

ailments. Honolulu Star-Bulletin, Honolulu

Kahng SE (2010) Ecological impacts of Carijoa 

riisei on black coral habitat. Final report to the 

Western Pacific Fisheries Management Council. 

Honolulu, Hawai‘i

Kahng SE, Benayahu Y, Wagner D, Rothe N (2008) 

Sexual reproduction in the invasive octocoral 

Carijoa riisei in Hawai‘i. B Mar Sci 82:1-17

Kahng SE, Grigg RW (2005) Impact of an alien 

octocoral, Carijoa riisei, on black corals in 

Hawai‘i. Coral Reefs 24:556-562

Kenyon J (1984) Black coral off Cozumel. Sea 

Frontiers 30:267-272

Lapian HFN, Barucca M, Bavestrello G, Biscotti 

MA, Bo M, Canapa A, Tazioli S, Olmo E (2007) 

A systematic study of some Black Corals 

species (Antipatharia, Hexacorallia) based on 

rDNA internal transcribed spacers sequences. 

Marine Biology 151:785-792

Moon HW, Song JI (2008) Taxonomy of the black 

coral family Myriopathidae (Anthozoa: 

Antipatharia) from Korea. Korean Journal of 

Systematic Zoology 24:251-263

NOAA (1983) 50 CFR 611 and 680. Foreign fishing 

and Western Pacific precious corals; 

implementation of fishing management plan. 

Federal Register 45

NOAA (2002) 50 CFR Part 660. Fisheries off West 

coast states and in the Western Pacific; 

precious corals fisheries; harvest quotas, 

definitions, size limits, gear restrictions, and 

bed classification. Federal Register 67

NOAA (2007) 50 CFR 665. Fisheries in the Western 

Pacific; precious corals fisheries. Federal 

Register 72:58259-58261

NOAA (2008) 50 CFR Part 665. Fisheries in the 

Western Pacific; precious corals fisheries; black 

coral quota and gold coral moratorium. Federal 

Register 73:47098-47100

NOAA (2012) 50 CFR Part 665. Western Pacific 

Fisheries; 2012 annual catch limits and 

accountability measures. Federal Register 

77:6019-6021

Noome C, Kristensen I (1976) Necessity of 

conservation of slow growing organisms like 

black coral. CCA Ecology Conference Bonaire 

11:76-77

Oishi FG (1990) Black coral harvesting and 

marketing activities in Hawai‘i - 1990. Division 

of Aquatic Resources, State of Hawai‘i

Olsen DA, Wood RS (1980) Investigations on 

black coral in Salt River Submarine Canyon St. 

Croix, U.S. Virgin Islands. FINAL SCIENTIFIC 

REPORT 80-12. Division of Fish and 

Wildlife U.S. Virgin Islands NULS-I Mission 

79-5 and 80-12

Opresko DM (2001) Revision of the Antipatharia 

(Cnidaria: Anthozoa). Part I. Establishment 

of a new family, Myriopathidae. Zoologische 

Mededelingen Leiden 75:343-370

Opresko DM (2002) Revision of the Antipatharia 

(Cnidaria: Anthozoa). Part II. Schizopathidae. 

Zoologische Mededelingen Leiden 76:411-442

Opresko DM (2003a) Redescription of Antipathes 

dichotoma Pallas, 1766 (Cnidaria: Anthozoa: 

Antipatharia). Zoologische Mededelingen, 

Leiden 77:481-493

Opresko DM (2003c) Revision of the Antipatharia 

(Cnidaria: Anthozoa). Part III. Cladopathidae. 

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(Cnidaria: Anthozoa). Part IV. Establishment 

of a new family, Aphanipathidae. Zoologische 

Mededelingen Leiden 78:209-240

Opresko DM (2005) New genera and species of 

antipatharian corals (Cnidaria: Anthozoa) from 

the North Pacific. Zoologische Mededelingen 

Leiden 79-2:129-165

Opresko DM (2006) Revision of the Antipatharia 

(Cnidaria: Anthozoa). Part V. Establishment 

of a new family, Stylopathidae. Zoologische 

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Opresko DM (2009) A new name for the Hawaiian 

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Antipathes dichotoma (Cnidaria: Anthozoa: 

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Opresko DM, Wagner D, Montgomery AD, Brugler 

MR (2012) Discovery of Aphanipathes verticillata 

(Cnidaria: Anthozoa: Antipatharia) in the 

Hawaiian Islands. Zootaxa 

3348:24-39

Padilla C, Lara M (2003) Banco Chinchorro: the 

last shelter for black coral in the Mexican 

Caribbean. B Mar Sci 73:197-202

Pallas PS (1766) Elenchus zoophytorum sistens 

generum adumbrationes generaliores et 

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cum selectis auctorum synonymis. HagaeComitum, Frankfurt am Main

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a review of reproduction of antipatharians 

(Cnidaria: Anthozoa: Hexacorallia). 

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Wagner D, Pochon X, Irwin L, Toonen RJ, Gates RD 

(2011c) Azooxanthellate? Most Hawaiian black 

corals contain Symbiodinium. Proceedings of the 

Royal Society B 278:1323-1328

Wagner D, Luck DG, Toonen RJ (2012a) The 

biology and ecology of black corals (Cnidaria: 

Anthozoa: Hexacorallia: Antipatharia). 

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Wagner D, Waller RG, Montgomery AD, Kelley 

CD, Toonen RJ (2012b) Sexual reproduction 

of the Hawaiian black coral Antipathes griggi 

(Cnidaria: Antipatharia). Coral Reefs 31:795-806

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Oceanology 3:56-61

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from Zhongsha Islands waters. Tropic 

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Kong waters with a description of a new 

species. Asian Marine Biology 1:101-105
         

       

           

     

Iridogorgia soft coral located off

the Northwest Hawaiian Islands.

Courtesy of the NOAA Office of

Ocean Exploration and Research.
                         

     

                     

           

                   

                     

                 

                   

                   

                 

                 

                   

                         

                   

                   

                     

                       

               

                     

         

                   

                   

           

                 

   

     

   

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

STATE OF DEEP-SEA 

CORAL AND SPONGE 

ECOSYSTEMS OF THE 

U.S. PACIFIC ISLANDS 

REGION 

Frank A. 

Parrish1*, Amy R. 

Baco2, 

Christopher 

Kelley3, and 

Henry Reiswig4

  

1 NOAA Ecosystem

Sciences Division, Pacific

Islands Fisheries Science

Center, Honolulu, HI  

* Corresponding author:

Frank.Parrish@noaa.gov

2Department of Earth,

Ocean, and Atmospheric

Science, Florida State

University, Tallahassee

3Department of

Oceanography, University

of Hawaii, Manoa, HI

4Department of Biology,

University of Victoria;  

Royal British Columbia

Museum, Canada

I. Introduction

The U.S. Pacific Islands Region consists of more than 50 oceanic

islands, including the Hawaiian Archipelago, the Commonwealth

of the Northern Mariana Islands (CNMI), and the territories of

Guam and American Samoa, as well as the Pacific Remote Islands

(Kingman Reef; Palmyra Atoll; Jarvis Island; Howland and Baker

Islands; Johnston Atoll; and Wake Island) and Rose Atoll. The

Pacific Island States in free association with the United States,

including the Republic of Palau, the Federated States of

Micronesia, and the Republic of the Marshall Islands, encompass

additional large portions of the central and western Pacific with

close ties to the U.S. 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; Fig. 1). Corals, sponges and other members of the

deepwater community are found in patchy distribution on these

oceanic ridges and have some of the highest levels of marine

endemism recorded anywhere on earth.

The 2007 State of Deep‐Sea Coral Ecosystems report (Parrish and

Baco 2007) detailed discovery and history of harvesting deep water

Corallium spp., Kulamanamana haumeaae (formerly Gerardia sp.

(Sinniger et al. 2013)), Isidella sp., the mesophotic black corals

Deep‐sea coral

community on a

Northwestern Hawaiian

seamount.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

Figure 1. Map of the Pacific Basin showing U.S. islands, their Exclusive Economic Zones (EEZ), and

Monument boundaries that comprise the Pacific Islands Region for the National Marine Fisheries Service. The

yellow is the area of Marine National Monuments and the red is the U.S. EEZ. At the atolls of Johnston,

Palmyra, and Wake the monument boundary is the same as the EEZ.

(antipatharians), and the associated

development of a survey and monitoring

program for the management of the “precious

coral” fishery.

The chapter listed the inventory of the known

structure forming corals and distinguished

these colonies from the reef building cold‐water

coral complexes found in the Atlantic and the

Gulf of Mexico. Also presented was

information on the distribution of corals in the

Hawaiian Archipelago and the absence of

survey effort in the wider expanse of the Pacific

Islands Region. As the harvesting and surveys

of corals relied on in‐situ visual observations

much was reported on the species associations

with deep‐sea coral communities including fish,

invertebrates, and even visitation by monk

seals. Aside from the limited activities of the

precious coral fishery, the list of potential

anthropogenic impacts to deep‐sea corals in the

region included impacts from placement of

undersea cables and cold seawater pipes, and

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Figure 2. Map showing submersible (Pisces IV, Pisces V) and ROV (RCV‐150) dive locations in the Hawaiian

Archipelago and Johnston Atoll where deepwater corals and sponges have been found. The yellow line is the

boundary of Marine National Monuments and the red is the U.S. EEZ. At Johnston Atoll the monument

boundary is the same as the EEZ.

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

the potential destructiveness of future cobalt‐

rich manganese (Mn) crust mining.

Until 2015, deep‐sea coral research in the Pacific

Islands Region was still focused almost entirely

in the Hawaiian Archipelago (Fig. 2). While

coral fragments from incidental dredging and

bycatch records indicates a wide distribution of

deep‐sea corals throughout the Pacific Island

Region, Hawai‘i is still the only location in the

region with assets to conduct such studies, and

even there the progress was sporadic,

dependent on when funding opportunities

present themselves through various requests

for proposals. Since the release of the 2007

national report the focus of deep‐sea coral

studies has shifted away from the intermediate

depth range (300‐500 m) where the commercial

harvest of precious corals motivated most of the

data collection, to coral/sponge communities

that are both shallower and deeper, along with

a greater emphasis on spatial modeling of

existing data. Roughly half of the publications

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

produced since the 2007 report are taxonomic in

nature. They include reviews of poorly

understood families, some new descriptions,

and some name changes. The rest of the papers

are a mix of growth, distribution and life

history studies. Work on the shallow water

black corals was the best represented segment

of the deep‐sea coral community including both

analysis of available data and some field

surveys. New information on black coral and its

history as a resource is presented in detail in

chapter 6. This follow‐up report differs from the

2007 report in that it includes information on

deepwater sponges in the region that was not

included in the first report.

II. Update on Scientific

Information

II.1. New Research – Overview

Since the 2007 status report (Parrish and Baco

2007), there have been a number of research

studies and publications on the region’s

deepwater corals. A complete list of new

publications is included at the end of this

chapter. Between 2007‐2014, there were a

number of deepwater coral and sponge field

projects concentrated in the Hawaiian

Archipelago, using the submersibles and

remotely‐operated vehicle (ROV) assets of the

Hawai‘i Undersea Research laboratory (HURL).

These projects involved 15 cruises funded

through HURL’s grant program and through

awards from the Papahānaumokuākea Marine

National Monument (PMNM), NOAA’s Center

for Sponsored Coastal Ocean Research

(CSCOR), Western Pacific Fishery Management

Council (WPFMC), and the National Marine

Fisheries Service (NMFS). Research topics

included precious coral age and growth,

habitat, distributions of coral and sponges, use

of corals for climate studies, studies of black

coral density and distribution, and effects of

invasive species on black coral populations.

There was also some shore‐based work

conducted using charter boats and three annual

(2008‐2010) research cruises using NOAA

vessels to support surveys of mesophotic coral

communities using remote cameras and mixed

gas divers that overlapped and contributed

new information to the status of shallow black

coral populations.

In addition to this fieldwork, analysis of data

from past expeditions and collections has

contributed to better understanding of deep‐sea

coral biology and ecology. Significant new

insights have been gained into the genetics

(Baco and Cairns 2012, Sinniger et al. 2013,

Figueroa and Baco 2014a,b), reproductive

biology (Waller and Baco 2007, Wagner et al.

2011c, Wagner et al. 2012), growth and age

(Parrish and Roark 2009, Roark et al. 2009), and

community structure (Long 2011, Long and

Baco 2014, Parrish 2015) of precious coral and

black coral species. Finally, international

collaborative efforts through the Census of

Marine Life Program, A Global Census of

Seamounts (CenSeam) have also provided

syntheses that provide a global context for the

Hawaiian Archipelago fauna. These include

analyses of global coral distributions and

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Figure 3. The three‐year CAPSTONE expeditions conducted mapping and deep‐sea surveys throughout the

U.S. Pacific Islands region.

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

habitat suitability modeling of corals on a

global basis (Rogers et al. 2007, Tittensor et al.

2009, Clark et al. 2011, Yesson et al. 2012,

Schlacher et al. 2014).

In 2015, NOAA initiated CAPSTONE: the

Campaign to Address Pacific monument

Science, Technology, and Ocean Needs. This

three‐year campaign brought together

resources from across NOAA to explore the

nation’s new central and western Pacific marine

national monuments and national marine

sanctuaries aboard NOAA Ship Okeanos

Explorer (Fig. 3). The campaign, led by NOAA’s

Office of Ocean Exploration and Research,

provided foundational science by discovering

and describing unknown deep‐sea habitats.

Science themes included the following:

 Acquiring data to support priority

monument and sanctuary science and

management needs;

 Identifying and mapping vulnerable marine

habitats – particularly high‐density deep‐

sea coral and sponge communities;

 Characterizing seamounts in and around

the Prime Crust Zone, the area of the Pacific

with the highest concentration of

commercially valuable deep‐sea minerals;

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

 Investigating the geologic history of Pacific

seamounts, including potential relevance to

plate tectonics and subduction zone biology

and geology; and

 Increasing understanding of deep‐sea

biogeographic patterns across the Central

and Western Pacific.

This three‐year campaign consisted of 23

cruises, 13 using the NOAA Deep Discoverer

ROV. Over 600,000 km2 of bottom was mapped

using multibeam, and 188 ROV dives explored

habitats between 240m – 6,000 m deep. More

than 360 samples of deep‐sea corals and

sponges were collected, a large proportion of

which are expected to represent new species.

These expeditions provided a first look at deep‐

sea coral and sponge habitats in U.S. Island

regions outside the Hawaiian Archipelago.

Among the most important initial discoveries

was the occurrence of numerous extremely

high‐density coral and sponge communities.

Initial information from the exploration is

available on NOAA’s Ocean Explorer website.

Scientific analysis of this rich trove of data and

information has just begun, and is not yet

reflected in following sections of this chapter.

II.2. Taxonomy and Species

Distributions

II.2.i – Corals

a. Coral taxonomy

Parrish and Baco (2007) reported 238 taxa of

deepwater corals from the Hawaiian

archipelago. The list included 75 taxa with

incomplete taxonomy. A total of 6 new genera

and 20 new species of octocorals,

antipatharians, and zoanthids have been

discovered in Hawai‘i since the 2007 report.

These are either new to science, or new records

for the Hawaiian Archipelago (Cairns and

Bayer 2008, Cairns 2009, Opresko 2009, Cairns

2010, Wagner et al. 2011a, Opresko et al. 2012,

Sinniger et al. 2013). Taxonomic revisions

currently underway for several groups of

corals, e.g., isidids, coralliids, plexaurids and

paragorgiids, are also likely to yield additional

species new to science and new records for

Hawai‘i.

Recent molecular phylogenetic and

morphologic studies of the family Coralliidae,

including Hawaiian precious corals, have

illuminated taxonomic relationships. These

studies synonymized Paracorallium into the

genus Corallium, and resurrected the genera

Hemicorallium (Ardila et al. 2012, Figueroa and

Baco 2014, Tu et al. 2015) and Pleurocorallium

(Figueroa and Baco 2014, Tu et al. 2015) for

several species, including several species in the

precious coral trade.

A molecular and morphological analysis of

octocoral‐associated zoanthids collected from

the deep slopes in the Hawaiian Archipelago

revealed the presence of at least five different

genera including the gold coral (Sinniger et al.

2013). This study describes the five new genera

and species and proposes a new genus and

species for the Hawaiian gold coral,

Kulamanamana haumeaae, an historically

important species harvested for the jewelry

trade and the only Hawaiian zoanthid that

appears to create its own skeleton.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

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Figure 4. Examples of structure‐forming deep corals found in the Hawaiian Archipelago. a) Pleurocorallium

secundum, b) Acanthogorgia sp., c) Iridogorgia magnispiralis, d) Antipathes grandis. Photo credit:

HURL.

b. Coral distribution and community

structure

Much of the funded work in the Hawaiian

Archipelago in the last decade has been related

to simply documenting species distributions. In

the 2007 status report, the Hawaiian coral beds

were divided into 2 zones, one at depths < ~600‐

700 m, and one > ~600‐700 m. The shallower

zone was dominated largely by precious corals,

but with a high diversity of other octocoral and

antipatharian species. The deeper zone was

dominated by a broad array of octocoral

families. Scleractinians were relatively rare at

all depths, and were primarily solitary (i.e. non‐

colonial) species. Records of colonial

scleractinians in the Archipelago continue to be

scarce. However, a large bed of dead

scleractinian Solenosmilia variabilis was

discovered in 2007 (Kelley unpubl. data), and

live corals were collected from a seamount in

the northwest end of the archipelago in 2016.

Baco et al. (2017) have reported deep reef

formations that appear to of constructed of this

species (see below).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

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From 2007 to 2014, little field work was directed

at deep‐sea corals (Fig. 4) and much of it was in

coral beds that had been documented years

prior. Consequently there was limited new

information on the spatial distribution of deep‐

sea coral in the Pacific Islands Region (Table 1).

Work includes cross contour surveys contracted

by the PMNM and climate‐related coral work

by universities that improved sampling on

seamounts in the central part of the

Northwestern Hawaiian Islands and conducted

the first exploration of the Necker Ridge which

extends south southeast from the central part of

the Hawaiian Archipelago (Morgan 2013,

Morgan et al. 2015). Habitat suitability

modeling studies as outlined below also

provide some insights onto potential locations

of deep‐sea coral abundance in the vastly

underexplored areas of this region.

Additional explorations as well as continued

analyses of pre‐2007 cruise video data and

sampling records, have provided more insights

into coral distributions and discoveries of new

species. A global synthesis of available data on

seamount coral taxa found that the Hawaiian

Archipelago is a diversity hotspot for deep‐sea

corals (Rogers et al. 2007).

The number of studies of the deep slopes of the

Hawaiian Archipelago is also beginning to

reach the critical mass needed to address

broader‐scale ecological questions in a more

quantitative way. The availability of the HURL

video archive as well as their video log

database have enabled these larger‐scale efforts.

The first of these efforts is a broad‐scale

synthesis of the HURL video log database,

encompassing the Northwestern Hawaiian

Islands and seamounts to the southeast of the

Archipelago that have been sampled with the

HURL submersible assets (Clark et al. 2011).

The goal of this study, funded by the

International Seabed Authority, was to compare

the megafaunal community structure of

seamounts with cobalt‐rich manganese (Mn)

crusts to those lacking such crusts, with the

southeast portion of the Hawaiian Archipelago

falling into the “cobalt‐rich” category. Over

30,000 records extracted from HURL’s database

from the Northwestern Hawaiian Islands,

Johnston Atoll, and Cross Seamount were

analyzed. No correlation was found with cobalt

or with feature type (island vs seamount, etc.),

instead depth seemed to be the overriding

factor related to the observed community

structure, with communities distinguished into

three depth groupings, those from dives with a

median depth of 227‐354 m, from dives with a

median depth of 357‐615 m and from dives

with a median depth of 745‐1799 m. Although

this analysis focused on all megafaunal

invertebrates, cnidarians made up about 30% of

the total records and were the key species

contributing to the differences among the 3

depth groups (Clark et al. 2011). Location was

also a significant factor, but was a secondary

factor to depth.

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175

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REG

  

Table 1. Primary structure‐forming deep‐sea corals and general attributes. The overall rating in the right column is a synthesis oNote: Hydrodendron gorgonoide is a chitinous hydroid, which while technically not a coral, nevertheless has similar character

Taxa ReefBuilding Abundance Maximum Colony Size Morphology Associations with other 

Invertebrates 

Colony SpatiaDisper

Acanella dispar No High Large branching many clum

Acanella weberi No High Medium branching clumAcanthogorgia sp. No High Small branching clum

Anthomuricea tenuispina No Medium Small branching clum

Antipathes sp. No Medium Large branching few bo

Aphanipathes sp. No High Medium branching few clum

Callogorgia gilberti No Medium Large branching bo

Calyptrophora sp. No Medium Large branching boChrysogorgia geniculata No High Small branching clum

Pleurocorallium secundum No High Medium branching few clum

Pleurocorallium niveum No High Small branching few clum

Coralliidae sp. (other) No High Medium branching few boEnallopsammia rostrata No High Medium branching clum

Enallopsammia cf. pusilla No High Medium branching clumEguchipsammia sp. No Medium Small branching clum

Hemicorallium laauense No High Medium branching few clumHemicorallium near laauense No Medium Large branching clum

Hydrodendron gorgonoidea No Medium Medium branching bo

Iridogorgia magnispiralis No High Large branching bo

Keratoisis sp. No High Large branching boKulamanamana haumeaae No High Large branching many clum

Leiopathes sp. No Low Large branching dispe

Lepidisis sp. No High Medium non‐branching bo

Madracis kauaiensis No Medium Small branching clumMadrepora oculata No Medium Medium branching dispe

Narella sp. No Medium Medium branching bo

Paracalyptrophora sp. No High Large branching clum

Paragorgia sp. No Medium Large branching clumVillogorgia sp. No Medium Medium branching clum

Plumarella circumoperculum No High Small branching clum

Solenosmilia variabilis Yes Medium Medium branching clum

Thouarella hilgendorfi No Medium Large branching clum

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

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Another recent study synthesized ROV video

from the Makapu’u precious coral bed on the

island of Oahu (Long 2011, Long and Baco

2014). The bed extends from a depth of

approximately 320 to 530 m. Despite the

relatively narrow depth range, depth was again

found to be the overriding factor correlated to

community structure, with other physical

parameters such as slope and rugosity also

playing a minor role. Intriguingly, although this

bed has always been referred to as a “precious

coral bed”, this quantitative assessment

indicated that the octocoral family Primnoidae

was actually the most abundant coral family

overall within the bed area. Coralliids did

dominate in one of the six depth bands, with

Corallium secundum (= Pleurocorallium secundum)

being the most abundant species there.

Interestingly, the peak in abundance of C.

secundum began about 400m, below the

maximum depth of previous harvesting

operations (Long 2011, Long and Baco 2014).

In contrast, surveys of deeper communities

(1400‐2000 m) on the Necker Ridge in 2011

revealed communities structured by factors

other than depth (Morgan 2013, Morgan et al.

2015). This study surveyed 6 sites along the

Ridge and found a significant difference in

communities on the northern side of the ridge,

which were dominated by octocorals,

compared to the southern side of the ridge,

which were dominated by crinoids. Diversity

also increased from south to north along the

ridge. A multivariate analysis of community

structure in relation to environmental factors

found that latitude, sediment cover, rugosity,

oxygen, salinity, and temperature, but not

depth, were most strongly correlated with

observed patterns.

Baco et al. (2017) recently reported the first

observations of deep‐sea scleractinian coral

reefs (bioherms) from the central North Pacific.

The reefs were found at depths of 535‐732 m on

six seamounts in the Northwestern Hawaiian

Islands and Emperor Seamounts, three of which

were within U.S. waters.

c. Coral population connectivity

While species lists and habitat suitability

provide first order insights into potential

connectivity of deep‐sea coral populations, the

real key to assessing connectivity is using

genetic methods. Genetic methods for assessing

connectivity are well developed in general, but

the paucity of sampling makes them

challenging to apply in the deep sea. The topic

is covered more broadly by Morrison et al. (this

volume). However, within the Hawaiian

Archipelago, phylogeographic and

phylogenetic work has focused primarily on the

precious corals and also on the primnoid

octocoral Narella. Hemicorallium laauense (Ardila

et al. (2012); Figueroa and Baco (2014) formerly

Corallium laauense) is the only Hawaiian deep‐

sea coral species that has published studies

using population genetics.

Data from three microsatellite loci in H. laauense

from 8 sites in the Main Hawaiian Islands and

southern end of the Northwestern Hawaiian

Islands indicated strong heterozygote

deficiency and no pattern of isolation by

distance (Baco and Shank 2005, Baco et al.

2006). The deepest site, at Kauai, in the middle

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

of the sampled locations, showed genetic

structure when compared to just about every

other site in the study. The Makapu’u bed on

the Island of Oahu also had some indication of

isolation for comparisons within the bed on the

scale of 1.6 km and for comparisons to most

other beds. These results indicate that the scale

of genetic structure within the Archipelago may

be very complex, and that the connectivity of

each feature or location should be assessed for

any management activities (Baco and Shank

2005).

Other types of genetic analyses such as DNA

barcoding and phylogeography may also yield

insights into connectivity, species distributions

and species ranges of Hawaiian corals. A study

designed to help develop barcoding markers

for octocorals found that species in the

primnoid genus Narella that occur in Hawai‘i

tend to have different haplotypes (DNA

sequences) within a single species that is found

at different depths. Of all the Hawaiian Narella

haplotypes, only one haplotype (from the far

Northwestern Hawaiian Islands) was also

found on Derickson Seamount, a deep

seamount south of the Aleutian Archipelago in

Alaska (Baco and Cairns 2012).

Taken together, this review of distributions of

Hawaiian corals from both genetics and species

lists, indicates that depth is a very strong factor

structuring deep‐sea coral communities,

potentially even over very small spatial scales.

Therefore depth structuring should be taken

into consideration in future management

efforts.

d. Modeling distribution and abundance

of corals

Although considerable effort has gone into

exploration and description of species to help

understand coral distributions in the Hawaiian

Archipelago, the continued high rate of species

discovery indicates that despite it being the best

studied Island group in the Pacific region, it too

is still drastically under‐sampled for deep‐sea

corals and other taxa. Several global syntheses

of coral distributions and habitat suitability

modeling included data from prior collections

in the Hawaiian Archipelago (Rogers et al.

2007, Tittensor et al. 2009, Tittensor et al. 2010,

Yesson et al. 2012). These provide a context for

results from Hawai‘i and also insights into the

likelihood of finding corals in the vast and

unexplored areas of the U.S. Pacific Islands

region (Tittensor et al. 2009, Davies and

Guinotte 2011, Yesson et al. 2012) and the

potential impacts of ocean acidification on

deep‐sea corals (Tittensor et al. 2010).

The first of these models relevant to corals was

by Tittensor et al. (2009) and focused on

scleractinians on seamounts. The results of this

modeling effort indicate very low habitat

suitability for scleractinians below 500 m across

the entire geographic range of U.S. territorial

waters in the Pacific Ocean. This work was later

expanded to other submarine features by

Davies and Guinotte (2011) and focused on five

of the most broadly distributed scleractinian

framework‐forming species (including

Enallopsammia rostrata and Madrepora oculata,

known to occur in Hawaii). They also

concluded that the North Pacific had very little  

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

suitable scleractinian habitat but a few features

that fall into U.S. waters in the southern

hemisphere did have a narrow depth band of

higher habitat suitability for certain species.

Yesson et al. (2012) focused on octocorals,

splitting them up into the seven major

suborders, and based on their model virtually

all of the sites within the U.S. Pacific Islands

region have high habitat suitability for

octocorals. The predicted lack of scleractinians

and high abundance of octocorals are consistent

with observations in this region (Baco 2007,

Parrish and Baco 2007), however a strong

caveat to these modeling efforts is that they are

done at relatively coarse geographic and

taxonomic resolution, and all of the models

remain to be ground‐truthed with in situ

explorations.

Besides providing insights into potential

distributions of corals, the models also provide

insights into the environmental parameters that

influence coral distributions. Recent modeling

efforts indicate the distributions of scleractinian

corals are related to depth, temperature,

aragonite saturation state, salinity, oxygen

concentration and saturation and dissolved

nutrients (Tittensor et al. 2009, Davies and

Guinotte 2011). Temperature, slope, salinity,

surface productivity, currents, oxygen and

calcite saturation state are correlated to the

distribution of octocorals (Bryan and Metaxas

2007, Yesson et al. 2012).

Habitat suitability modeling for precious corals

in the Main Hawaiian Islands was funded by

NOAA’s Deep Sea Coral Research and

Technology Program. The project “Predictive

Modeling of the Distribution of Deep‐Sea

Corals in the Main Hawaiian Islands” (PIs John

Rooney, Amy Baco and Michael Parke) used

data from the Main Hawaiian Islands in

attempt to identify suitable habitat of Corallium

secundum (pink coral; =Pleurocorallium

secundum) and K. haumeaae (gold coral), that are

important targets in the precious coral fishery.

A total of 1227 observations of these taxa were

made in the Main Hawaiian Islands during 102

of the more than 1100 submersible dives

completed in the region by the Hawai‘i

Undersea Research Laboratory. An analysis of

circulation data from ocean general circulation

models and tidal models indicates that in this

region, at depths below the main thermocline

(300 to 500 m), tidal velocities are an order of

magnitude larger than the general circulation

flows. Deep‐sea coral growth is thought to be

enhanced in areas with higher current velocities

that would increase the availability of food.

Accordingly, maximum baroclinic tidal

velocities at a depth of 400 m were calculated

using output from a tide model developed by

Carter et al. (2008) for a portion of the Main

Hawaiian Islands on a one by one kilometer

grid (Fig. 5). A high resolution grid of all

available bathymetry data was also obtained

from the Hawai‘i Mapping Research Group at

the University of Hawai‘i. The modeling effort

was unable to identify any significant

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

Figure 5. Maximum modeled current speeds at depth. The red end of the color spectrum indicates stronger

currents.

predictions of coral occurrence but based on the

observations it is hypothesized that these

species of deep‐sea corals prefer areas with

current velocities 0.5 ‐ 0.85 m/s

(https://deepseacoraldata.noaa.gov/other‐

studies/fy10‐08).

In a subsequent study, Bauer et al. (2016)

modeled habitat suitability for 16 deep‐sea

coral groups, based on ecological importance,

management considerations, and the number of

presence records. Models included Corallium

and the gold coral K. haumeaae. The Corallium

model was among the best fits, with distance to

shore, total curvature at 5 km scale, and annual

bottom temperature as the most important

predictor variables.

II.2.ii – Sponges

a. Sponge taxonomy and distribution

Sponges in the phylum Porifera are presently

divided into 4 classes: Calcarea, Demospongiae,

Hexactinellida, and the “revived” class

Homoscleromorpha, which until recently has

been considered to be an order of demosponges

(Gazave et al. 2010). In general, most sponges in

depths above 50 m are either demosponges,

calcareans, or homoscleromorphans, whereas

most sponges below 400 m are hexactinellids

(Reiswig unpub. data, HURL database).

Deepwater sponges in the Pacific Islands region

are more poorly known than the deepwater

corals and only those large enough to be

considered structure‐forming are included in

this report (Table 2, Fig. 6). The major

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

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transition between the hexactinellids and the

other classes of sponges in this region appears

to be below 200 m, perhaps as a result of higher

water temperatures and light penetration above

that depth (HURL database). However, this

could also be a result of under‐sampling within

the 50‐200 m depth zone, which for the most

part falls between the routine operating depths

of SCUBA and deepwater vehicles such as

submersible and ROVs. For simplicity, this first

inclusion of deepwater sponges in the report

for the Pacific Islands Region will focus on

species found at depths below 200 m, and is

largely derived from the HURL on‐line

Deepwater Animal Guide

(http://www.soest.hawaii.edu/HURL/) and

related database records (Fig. 6).

Table 2. Structure‐forming attributes of deep‐sea sponges in Hawaii. The overall rating in the right column is a

synthesis of the rest of the attributes.

Taxa ReefBuilding Abundance Colony 

Size 

Colony 

Shape 

Spatial 

Dispersion 

Overall 

Rating 

Bolosoma sp. (all) no medium large pedunculate sphere both medium

Farrea near occa no high medium anastomosing clumped medium

tubes

Poliopogon sp. (all) no high large hemispherical clumped high

Regadrella sp. 1 no medium medium tube‐like clumped medium

Saccocalyx cf. no medium large pedunculate cup dispersed medium

pedunculatus

Semperella sp. (all) no high medium columnar clumped high

Sericolophus no high medium pedunculate spoon clumped high

hawaiicus

Tretopleura sp. 1B no medium large elongated plate both medium

and sp. 2

Walteria sp. (all) no medium medium tube‐like both medium

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B

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

Figure 6. Examples of structure forming deep sponges found in the Hawaiian Archipelago: a) Sericolophus

hawaiicus, b) Poliopogon sp4, c) Poliopogon sp1, d) Bolosoma sp2. Photo credit: HURL.

a.(i) Calcarea and Demospongiae below 

200 m 

A single colony identified from a photograph as

a calcarean species in the genus Clathrina, has

been included in the HURL database and online

identification guide for many years. In 2017,

this rare species was encountered and collected

during an Okeanos Explorer cruise to the

Johnston Atoll monument area. Microscopic

examination determined that it was actually a

misidentified and potentially new species of

hexactinellid in the family Tretodictyidae.

Therefore, at the present time, there are no

confirmed published or unpublished accounts

of deepwater calcareans being found in the

Pacific Islands region.

Chave and Malahoff (1998) listed seven species

presumed to be demosponges that have been

photographed from submersibles operated by

HURL. One of these has subsequently been

collected and identified as a species of

hexactinellid. Another of these, Corallistes sp.,

has been only identified from photographs but

is similar to confirmed species so is listed in the

appendix as cf. Corallistes sp. The remaining

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a.(ii) Hexactinellida below 200 m 

             

             

                 

       

   

           

   

     

   

           

           

             

           

         

   

               

           

             

                 

             

           

                     

               

               

               

                 

             

       

                 

     

             

               

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

five species were described with only

descriptive names so are not included.

The HURL database has 9 species of

demosponges including cf. Corallistes sp. that

occur below 200m in Hawai‘i. Of these, 5 were

collected and identified by spicule examination:

Penares sp., Poecillastra sp. 1, Poecillastra sp. 2,

Hamacantha (Vomerula) n. sp. and Stelodoryx sp.

The remaining 4, cf. Corallistes sp., cf. Petrosia

sp., cf. Polymastia sp. 1 and cf. Polymastia sp. 2

were identified from images only but all show

clear similarities to confirmed species in those

genera. During the preparation of this report,

additional demosponge specimens were

collected during HURL submersible and Deep

Discoverer ROV dives that have not as yet been

entered into the HURL or National Deep‐Sea

Coral and Sponge databases. These include a

Geodia sp., an unidentified species in the order

Haplosclerida, another Poecillastra sp., an

unidentified species of Characella, an

unidentified species of Desmacella, a

dendoricellid in the genus Pyloderma, and

several species in the family Cladorhizidae, .

All were identified from spicules or spicule

images by Dr. Rob van Soest in the

Netherlands. Of these 18 total species, cf.

Corallistes sp., Penares sp., Poecillastra sp. 2,

Polymastia sp. 1, Characella sp., Stelodoryx sp.

Pyloderma sp. were found in discrete beds that

may have formed from localized settlement

events. While several of these can found in

moderate to high abundance, the colony sizes

of these sponges are relatively small and few if

any have been observed with associated fauna.

None are therefore considered to be major

structure forming species, however it is worth

noting here that based on these new collections,

there may be a greater prevalence of

demosponges below 200 m than previously

thought.

Prior to 2015, 118 hexactinellid specimens had

been collected in Hawaiian waters dating back

to the Albatross expedition in 1902. Three of the

five currently recognized orders

(Amphidiscosida, Lyssacinosida, and

Sceptrulophora) and seven of the currently

recognized families (Pheronematidae,

Uncinateridae, Euretidae, Farreidae,

Tretodictyidae, Euplectellidae, and Rossellidae)

are represented in these collections. At least 49

distinctly different taxa are among the

specimens. However, to date, only three have

been identified to the species level: Sericolophus

hawaiicus Tabachnick and Levi 2000, Walteria

flemmingi Schultze 1886, and Farrea occa

Bowerbank 1862. Seven types are listed as being

“near” a previously described species because

spicule examination has shown they are close to

but not an exact match and therefore may be

different. The remaining 42 types are believed

to be potentially new, undescribed species,

which are the most of any region in the U.S. or

for that matter, in the world (Reiswig unpub.

data). A collaborative effort that includes two of

the authors of this report (Kelley and Reiswig)

in addition to Dr. Craig Young and Dr. Martin

Dohrmann is now underway to publish species

descriptions for this somewhat “neglected”

fauna. In addition to the collections, at least 8

additional distinctly different hexactinellids

have been captured on video during HURL

dives and are included in this report’s online

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species list but have not as yet been collected.

At the preparation of this report, there are

believed to be at least 57 species of

hexactinellids in Hawaiian waters that have

been recorded from 330 m to over 2,000 m.

However, recent CAPSTONE collections (that

included from waters deeper than 2,000 m, as

well as from U.S. Pacific Island areas outside

the Hawaiian Archipelago) will significantly

increase these numbers once the specimens

have been carefully examined.

1.0 Order Amphidiscosida

1.1 Family Hyalonematidae  

Chave and Jones (1991) and Chave and

Malahoff (1998) reported the presence of two

families of Amphidiscosida in Hawai‘i:

Hyalonematidae and Pheronematidae.

However, further investigation determined

their unidentified species of Hyalonema was

actually a pheronematid that was subsequently

named Sericolophus hawaiicus (Tabachnick and

Levi 2000). Two species of hyalonematids were

recently collected during CAPSTONE dives

around Hawai‘i. Therefore even though the

initial report of this family being in Hawai‘i

was found to be incorrect, its presence has now

been confirmed.  

1.2 Family Pheronematidae

All other Hawaiian amphidiscosidans

identified to date are pheronematids, with

representatives from three genera: Poliopogon,

Semperella, and Sericolophus. Pheronematids

have been both collected and recorded on video

between 338 m and the 2000 m maximum

operating depth of the Pisces submersibles. This

group no doubt extends much deeper. While

most species are found attached to hard rocky

substrates, S. hawaiicus, the shallowest species,

is found exclusively on soft substrates. S.

hawaiicus has an elongate tuft that both anchors

the sponge into the sediment as well as serving

as a stalk (Fig. 6a). This species is considered to

be structure forming because it is typically

found in very high density beds and with a

variety of commensals since it often provides

the only firm elevated surfaces in the area.

Some of the other pheronematids are both large

and also form beds with densities close to 1/m2.

Large lateral oscula in many of these species

attract commensal crabs and ophiuroids that

appear to use them for shelter. Until the

taxonomy of family is completely worked out,

all species of pheronematids in Hawai‘i should

be considered as structure forming due to both

their size and abundance (Fig 6b,c).

2.0 Order Sceptrulophora

The order Sceptrulophora (formerly

Hexactinosida, in part) is represented in this

region by four families: Euretidae, Farreidae,

Tretodictyidae, and Uncinateridae.

2.1 Family Euretidae

Three different species of euretids have been

identified from specimens collected during

HURL dives in Hawai‘i. The latest of these, a

suspected new species of Lefroyella, was

collected in 2013 and like its New Zealand

congener, Lefroyella decora, was found to have a

large number of commensal anemones growing

through its framework. These anemones are

believed to be in the family Edwardsiidae

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(Fautin, pers comm.), but this has not as yet

been confirmed. Additional euretids were

collected during CAPSTONE dives bringing the

total number of species tentatively identified as

being euretids to 9. However, these are all

relatively uncommon and therefore are not

considered major structure forming sponges in

this region.  

2.2 Family Farreidae

All of the farreid specimens collected during

HURL submersible dives around Hawai‘i have

similarities to Farrea occa, which has been

tentatively identified as being in this region

from a single specimen. These species are

therefore described as Farrea near occa or Farrea

near occa erecta based on whether their colony is

bushy (the former) or taller and more upright

(the latter). Farreids have been recorded from

350‐2400 m in Hawai‘i and are relatively

common. In some locations F. near occa, but not

F. near occa erecta, can be found in beds of

modest densities. Another interesting

distinction between the two types is the

presence of a commensal antipatharian found

growing on F. near occa but not on F. near occa

erecta. This antipatharian appears to be a new

species and is presently under investigation.

Other colonies of F. near occa have been seen

completely covered with zoanthids which again

has not been observed with colonies of F. near

occa erecta. Collections of farreids during

CAPSTONE have revealed the presence of at

least 2 other genera around Hawai‘i:

Aspidoscopulia and Lonchiphora as well as a

possible third new genus that presently doesnʹt

match any of the known genera in this family.

Aspidoscopulia colonies are large and can be

found in relatively dense beds. However, until

these new specimens can be examined more

carefully, F. near occa will be listed here as the

only structure forming farreid in this region.  

2.3 Family Tretodictyidae

Although two species in this family are listed in

this report’s online species list, only one has

been positively identified in this region and has

been assigned to the genus Tretodictyum. The

specimen is vase‐shaped and was collected on

hard substrate at a depth of 687 m. This species

is neither large nor abundant and therefore is

not considered to be structure forming. As

mentioned previously, a potentially new

species of branching tretodictyid was collected

several months ago, which had been previously

identified from photographs as a calcarean.

Colonies were moderately abundant at the

CAPSTONE dive site and were furthermore

found to have large numbers of what appeared

to be commensal zoanthids. This is likely a new

species however that determination requires

additional investigation.

2.4 Family Uncinateridae

The presence of this order in Hawai‘i was

unknown until 2011 when the identification of

four specimens collected in 2003 were assigned

with several other specimens collected in 2007

and 2009 to the genus Tretopleura in the family

Uncinateridae. Three additional specimens do

not fit into any known genera and are therefore

listed as “Uncinateridae new genus new

species.” All of the species in this family attach

to hard substrates at depths of 685‐1850 m.

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Commensal crabs and ophiuroids are typically

observed on the Tretopleura colonies, which can

reach over a meter in height. Tretopleura sp. 1B

and sp. 2 are relatively common and are

therefore considered structure forming.

3.0 Order Lyssacinosida

Lyssacinosidans are relatively common in this

region of which all have been assigned to one of

two families: Euplectellidae and Rossellidae.

3.1 Family Euplectellidae

Euplectellids are the most diverse family of

hexactinellids in Hawai‘i with 18 potential

species having been identified from specimens

and video images. Thirteen genera have been

identified to date and include one in the

lophophytous subfamily Euplectellinae, four in

the pedunculate subfamily Bolosominae and 8

in the vase‐like and tube‐like subfamily

Corbitellinae. While Dictyocalyx gracilis was

reported to have been collected near Hawai‘i

(Tabachnick 2002), the 2 specimens of this

genus actually collected in Hawai‘i are not the

same and are believed to be a new species.

Euplectellids are found from 330 m to over

4,000 m, with those in the genus Regadrella

being observed in the shallowest depths while

pedunculate euplectellids only appearing at

depths below 500 m. All species attach to hard

substrates and several have been observed in

aggregations. Some species in both the

Bolosominae and Corbitellinae can grow to

relatively large sizes (Fig. 6d). Most

euplectellids are inhabited by commensals such

as ophiuroids and crustaceans. In some species  

with small or covered primary oscula, early

colonization leads to commensal shrimp

becoming “trapped for life” inside their atrial

cavities (Baba 1983). For the purpose of this

report, several species of euplectellids found in

the genera Bolosoma, Saccocalyx, and Regadrella,

and Walteria are considered structure forming in

this region.  

3.2 Family Rossellidae

Until CAPSTONE, rossellids were represented

in Hawai‘i by 9 species, 7 of which have been

placed into two genera: Bathydorus and

Caulophacus. The two species of Bathydorus are

vase‐like, with the specimen of Bathydorus near

laevis being a huge caldron sized sponge while

the specimen of Bathydorus sp. was not. These

two species, along with other similar looking

vase sponges identified from video as only

rossellids, have been recorded at depths of 350‐

1760 m. The five species of Caulophacus are

pedunculate with small bodies supported

above the substrate by a relatively long stalk.

Both pedunculate and non‐pedunculate

rossellids can be found in small aggregations

but none are considered structure forming in

this region. CAPSTONE collections of rossellids

around Hawai‘i included members in

additional genera of both pedunculate and non‐

pedunculate species. The most noteworthy of

these was a specimen from an extremely large

colony of an undescribed genus in the

subfamily Lanuginellinae. While this colony

had numerous associated fauna, the species is

rare and therefore is therefore not considered to

be structure forming at this time.  

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Figure 7. Maps of the Hawaiian Archipelago and Johnston Atoll showing the locations where deepwater

octocorals (purple, upper left), scleractinians (orange, upper right), antipatharians (green, lower left), and sponges

(blue, lower right) have been found. For additional detail see maps in Appendix. The yellow line indicates the

monument boundaries and the red line is the EEZ boundary. The two are the same around Johnston Atoll.

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II.3. Coral Growth, Biology and

Reproduction

The Western Pacific Fishery Management

Council placed a moratorium (2007) on the

harvesting of gold coral (K. haumeaae) (Fig. 8)

for the precious coral fishery until a validation

study of conflicting growth estimates could be

conducted. Gold coral colonies that were

measured and marked up to 9 years earlier

were revisited and re‐measured (Parrish and

Roark 2009). The measured change in growth

was statistically indistinguishable from zero,

indicating that gold coral growth is much

slower than the annual growth estimate

derived from basal stem ring‐counts previously

relied on for the fishery. The duration of the re‐

measurement period was too short to

effectively evaluate the validity of longer

lifespan estimates from radiocarbon dating that

yielded a linear growth rate of 2.2 ± 0.2 mm yr‐1

but clearly indicated that gold coral growth was

very slow. The main criticism of using

radiocarbon measurements to determine the

lifespan of K. haumeaae and Leiopathes sp.

(estimated at up to 2,742 years and 4,265 years,

respectively) was that the corals might be

feeding on old dead carbon thereby giving

unrealistically old ages. By reproducing the

radiocarbon bomb curve in a Hawaiian K.

haumeaae specimen (Roark et al. 2009) and a

Gulf of Mexico Leiopathes sp. specimen (Prouty

et al. 2011) these studies clearly demonstrate

that these deep‐sea corals are feeding on

recently exported young fresh particulate

organic carbon from surface waters clearly

demonstrating the validity of the extreme

longevity and slow radial growth for both

species (see Prouty et al., this volume, on age,

growth rates, and paleoclimate studies). U‐

Series dating of stony coral Enallopsammia

rostrata specimens from 480 to 788 m water

depth in the Line Islands indicate life spans

ranging from 209 ±8 to 605 ±7 years, radial

growth rates ranging from 0.012 to 0.073 mm yr‐

1 and with linear growth rates ranging from 0.6

to 1.9 mm yr ‐1 (Houlbrèque et al. 2010). These

Figure 8. The purple stoloniferan Clavularia

grandiflora growing on a Kulamanamana

haumeaae colony. Photo credit: HURL.

Figure 9. The invasive Carijoa riisei growing

on shallow water black coral. Photo credit:

HURL.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

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results show E. rostrata is relatively slow

growing deep‐sea coral.  

Recent reproductive studies on Hawaiian deep‐

sea corals includes work on the morphological

structures of precious gold coral (K. haumeaae),

pink coral (Pleurocorallium secundum), and red

coral (Hemicorallium laauense) (Waller and Baco

2007). All 3 species appear to be gonochoric

broadcast spawners. The reproductive biology

of the shallow water black corals (including

Antipathes grandis, Antipathes griggi, Cirrhipathes

cf. anguina, Stichopathes echinulata, Stichopathes

sp., Aphanipathes verticillata), and deepwater

corals (Leiopathes sp. and Alternatipathes alternata

[=Bathypathes alternata]) (Wagner et al. 2011c,

Wagner et al. 2012) has also been examined.  

Wagner et al. (2011b) found that 10 out of 14

Hawaiian black coral species collected between

10 and 396 m contained the symbiotic algae

Symbiodinium and thus perhaps should not be

considered strictly azooxanthellate. Histological

examination of five species showed low

densities of Symbiodinium‐like cells inside

gastrodermal tissues. The authors concluded

that the Symbiodinium were unlikely to play a

significant role in the nutrition of the black

corals.  

II.4. Species Associations with

Deep-Sea Corals and Sponges

The 2007 report summarized information on

species associations with deep‐sea corals and

highlighted concerns over the colonization of

shallow water black coral by the parasitic

octocoral Carijoa sp. (originally identified as

Carijoa riisei; Fig. 9). Wagner et al. (this volume)

discuss the improved understanding of this

parasitic association. Many additional

associations of invertebrate species with deep‐

sea corals and particularly with sponge

communities (Fig. 10) have been documented.

The exceptional imagery collected during the

CAPSTONE cruises have expanded the number

of observed associations and along with

collected samples will be a future area of

research focus.

II.5. Research Priorities

The priorities for future work in the region on

both deep‐sea corals and sponges remain

largely the same as those identified for deep‐sea

corals in the 2007 report, and can be divided

into: 1. Exploration and mapping, 2. Taxonomy

and connectivity, 3. Biology, ecology and

environmental parameters, and 4. Resilience of

deep‐sea coral communities. In 2014, NOAA

brought together over 20 researchers and

resource managers to identify and prioritize

information needs to increase our

understanding of deep‐sea coral and sponge

ecosystems in the U.S. Pacific Island Region.

The workshop participants identified priority

research questions and specific information

needs and activities (Parrish et al. 2014). The

results of the workshop informed NOAA’s

Deep‐Sea Coral and Sponge Fieldwork

Initiative and the CAPSTONE expeditions, and

are reflected in the priorities below:

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

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Figure 10. Examples of invertebrates found associated with Hawaiian deep‐sea corals (a‐ophiuroid, b‐crinoid, c‐

squat lobster, d‐asteroid feeding on a bamboo coral; Photo credit: HURL) and sponges (e‐squat lobster, f‐shrimp

and ophiuroids, g‐antipatharian growing out of sponge, h‐ophiuroids; Photo credit: NOAA).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

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1. Exploration and mapping

Prior to 2015, there had been little or no

exploration of the U.S. Pacific Islands Regions

except for the Hawaiian Archipelago. There is a

need to expand surveys to under sampled

regions in the U.S. Pacific including the

Marianas, American Samoa and the Pacific

Remote Islands Marine National Monument.

Exploring the rest of the U.S. Pacific is the

highest priority for the region and represents a

real challenge. Applying consistent taxonomy,

and where possible, collecting genetic samples

in order to evaluate the degree to which various

assemblages are connected, will begin to

elucidate biogeographic patterns at the Pacific

basin scale. Even within the Hawaiian

Archipelago, where research tools are available,

most information on the distribution of deep‐

sea corals and sponges was restricted to depths

shallower that 2000 m and little was known

about the deep‐sea fauna found below this

depth.

2. Taxonomy and connectivity

Taxonomy using both traditional

morphological methods and recently developed

genetic methods continues to be a high priority

for all deep‐sea coral and sponge work because

so many of the specimens are new to science.

While gene flow in shallow water ecosystems is

relatively well‐understood, the gene flow in

deep‐sea coral ecosystems is poorly understood

and their role in connectivity is an obvious

emphasis for future work in the U.S. Pacific

Islands region. CAPSTONE collected numerous

samples of corals and sponges (including

genetic sampling) that have already begun to

identify new species (e.g., Cairns 2017),

however, population‐level studies were

constrained by limited sample sizes.

3. Biology, ecology, and environmental

parameters

Much of the basic biology and ecology of major

structure‐forming deep‐sea corals and sponges

in the region are poorly known, as are the

environmental variables that govern where

deep‐sea coral and sponge communities

colonize and develop into mature assemblages.

Having information on the conditions needed

to support these ecosystems is valuable, as this

provides the parameters needed for planned

modeling activities and contributes to the

growing body of climate research. To date there

are no published habitat suitability models for

deep‐sea sponges. The Pacific Islands, with its

large protected areas in the form of marine

national monuments, is an ideal place to look

for environmental change using data loggers at

established study sites.

4. Resilience of deep‐sea coral communities

It is not known how long it takes for deep‐sea

coral or sponge individuals or communities to

recover following disturbance. Deep‐sea corals

and probably many sponges can grow

extremely slowly and are vulnerable to impacts

from activities that disturb the seafloor.

Additionally, ocean acidification may affect

corals’ ability to grow and maintain their

structures. There is a need to understand

community development that considers overall

natural recruitment, growth, and mortality over

large spatial scales.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

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Opportunities for addressing these research

priorities need to take into account the loss of

research and monitoring assets in the region. In

2013 NOAA discontinued funding for the

Hawai‘i Undersea Research Laboratory, which

operates the Pisces 4 and Pisces 5 submersibles,

ROV, and support vessel. These have been the

key and nearly exclusive assets for in situ

explorations in the U.S. Pacific Islands Region.

Loss of these assets will severely impact deep‐

sea coral and sponge work in Hawai‘i and the

U.S. Pacific Islands Region. The availability of

other similar assets typically requires that

highly competitive proposals based on

hypothesis‐driven research be submitted to

funding entities. The success of CAPSTONE

highlights the value of exploratory work,

however it may be incredibly challenging to

bring submersible and ROV assets here on a

sustained basis in the future.

III. Update on Management

of Deep-Sea Coral and

Sponge Habitats

III.1. New Information on Impacts

and Stressors

III.1.i – Precious coral fishery

Hawai’i has had the only significant black and

precious coral fisheries in the U.S. There have

been no reported commercial catch reports for

precious coral since the 2007 report. Harvest of

the precious coral (e.g., Corallium spp.,

Hemicorallium spp., Pleurocorallium spp, K.

haumeaae) has been dormant since 2000

although a permit was obtained in 2012 for

harvesting in the Makapu’u bed using a small

ROV system but is as yet unused. In 2011 sales

reports of shallow water black coral were filed

but cannot be described in detail because the

number of participants is too small to report

publicly due to confidentiality requirements.

Robin Lee, long time black coral diver and key

participant in the fishery passed away in 2012,

and recently submitted landing logs indicate

new participants have joined the fishery.

III.1.ii – Undersea cables

The laying of undersea cables in the channels of

the Hawaiian Islands to deliver power from

planned wind and solar farms on outer islands

(e.g., Molokai, Lanai) is the region’s most

immediate potential impact concern. Most of

the known shallow‐water black coral

population is found in these channel waters

and are the focus of harvesting by divers. Tracts

of zooxanthellate scleractinian corals (e.g.,

Montipora capitata, Leptoseris hawaiiensis) also

occur at the mesophotic depths between 30‐150

m and are fragile, growing on the slopes and

the submerged land bridge that connects the

islands and could incur some damage if cables

are laid on them. Azooxanthellate corals in the

deeper channels between Oahu and Molokai

are also at risk of damage from cables.

However, at the moment, the state has taken

the effort to survey potential cable routes and

has already diverted proposed routes around

known coral beds such as that off Makapu’u.

III.1.iii – Undersea mining

The interest in mining cobalt‐manganese crust

from the seafloor between 400‐4000 m in the

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U.S. Pacific was identified in the 2007 report

and continues to be discussed as an option for

the future. Although this is unlikely to occur in

areas such as the Hawaiian Archipelago due to

its young age and the fact that much of the

archipelago has been designated as a marine

national monument, other, older island groups

in the U.S. Pacific may be more likely targets.

However, Cretaceous seamounts (i.e., 60‐120

ma) with presumed extensive Mn crusts do

exist in the vicinity of the archipelago and

could be targeted by the industry in the future.

There is enough concern about the impacts of

mining that the International Seabed Authority

(ISA) funded data syntheses to provide the first

ever comprehensive description of taxa present

on cobalt‐rich crusts (see Clark et al. (2011) and

Schlacher et al. (2014)). Additional studies of

this kind are clearly warranted, particularly in

light of the fact that 3 exploratory leases have

already been issued by the ISA on prospective

Mn crust mining sites in the Western Pacific.

III.2. New or Planned

Management Actions

III.2.i – Precious coral fishery

The only change in federal regulation for the

precious coral fishery is a 5‐year moratorium on

the harvest of live gold coral due to its reported

slow growth rate (Roark et al. 2006). A

subsequent study (Parrish and Roark 2009)

using mark and re‐measurement techniques

reported similar findings and the moratorium

was extended until 2018. Other proposed

actions relevant to the coral fishery included a

proposal in 2009 to list the genera Corallium and

Paracorallium under the Convention on

International Trade in Endangered Species of

Fauna and Flora (CITES) Appendix 2 but this

proposal did not pass.

III.2.ii – Designation of new Marine

National Monuments

In January 2009, Presidential proclamations

8335‐8337 designated waters off CNMI as the

Marianas Trench Marine National Monument;

Kingman Reef, Palmyra Atoll, Jarvis, Howland,

Baker, Johnston and Wake Islands were

designated the Pacific Remote Islands Marine

National Monument; and Rose Atoll was made

the Rose Atoll Marine National Monument

under the Antiquities Act. These new

Monuments added to the existing

Papahānaumokuākea National Marine

Monument (est. 2006) to form the largest

network of marine monuments in the U.S. Their

mandate is to protect the unique natural and

cultural resources within the region through the

advancement of scientific research, exploration

and public education. Combined this area

encompassed a total 800,076 km2 with each

marine national monument extending seaward

50 nm from mean low water the seabed. They

are co‐managed between the Department of

Interior and the Department of Commerce in

coordination with federal partners (e.g.,

Department of Defense) and the

State/Territorial governments. No commercial

fishing is allowed in any of the monuments,

which precludes the expansion of the precious

coral fishery into these areas.

In 2014, the boundaries of the Remote Pacific

Islands National Marine Monument around

Jarvis, Wake and Johnston Atoll were expanded

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

out to the 200 nautical mile U.S. EEZ by

Presidential Proclamation 9173. The newly

protected areas contain additional seamounts

that have since been confirmed to include

important deep‐sea coral and sponge resources.

The monument now encompasses an area of

1,057,494 km2 (308,316 square nautical miles).

These virtually unexplored areas were the

target of mapping and ROV surveys by NOAA

ship Okeanos Explorer under CAPSTONE.

The Papahānaumokuākea Marine National

Monument management plan was finalized in

2008, and in 2010 the area was inscribed as a

natural and cultural World Heritage Site. The

Monument was expanded by Presidential

Proclamation 9478 to 1,508,870 km2 (582,578

square miles), making it the largest protected

area ‐ terrestrial or marine ‐ on the planet.

III.3. New Priorities for

Management

III.3.i – Alternative Energy

Hawai‘i’s push for alternative energy has

prompted a number of feasibility studies

including some initial surveys of potential

cable‐laying routes. Management groups

involved in these plans are studying the

potential impacts and examining the best

alternatives. An example is the Hawaiian

Islands National Humpback Whale Sanctuary,

which spans the channel waters of the Main

Hawaiian Islands, partnering with NOAA

scientists and the University of Hawai’i to

spatially model data from their studies on

mesophotic coral ecosystems in the Au’au

channel between the islands of Maui and Lanai

(Costa et al. 2012). The Bureau of Ocean Energy

Management also supported modeling of

shallow, mesophotic and deep‐sea corals (Bauer

et al. 2016) in order to inform potential future

alternative energy development. There has also

been an interagency meeting to look at

potential impacts of implementing Ocean

Thermal Energy Conversion (OTEC) using

intake pipes laid down slope 800‐1000 m to

bring deep cold water to the surface for energy

production.  

III.3.ii – Ocean cage aquaculture

Limited offshore cage aquaculture of fishes has

occurred on the leeward coast of a couple of the

Main Hawaiian Islands and there is interest in

expanding this activity, which could involve

use of mooring anchors potentially at deeper

depths. It is not known if this type of

development would impact deep‐sea coral

communities significantly.

IV. Conclusions

In the first seven years since the 2007 report,

deep‐sea coral research focused on using

available data from Hawai‘i to study the

biology, reproduction and growth of select

coral taxa – especially antipatharians. Notable

advances included taxonomy, preliminary

habitat suitability models for deepwater corals,

and the first steps in the characterization of the

sponge community. Hawai‘i’s deep‐sea

sponges are far less well studied than are the

deep‐sea corals. Much of the U.S. Pacific Islands

has been recently designated as new Marine

National Monuments and yet still little or no

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

information had been collected on the deep‐sea

coral and sponge communities in areas outside

of the Hawaiian Archipelago. The monument

designations will limit expansion of domestic

commercial harvesting especially for precious

coral resources that are protected as part of the

seafloor. However, much of this area is remote

making it difficult to monitor the degree of

illegal fishing by foreign fleets. Other potential

impacts include cable‐deployment and

development of oceanic cage aquaculture, and

for the adjacent high seas region, deep‐sea

mining. Finally, the region’s ability to continue

and expand studies of deep‐sea corals in

support of management using the submersible

and ROV assets operated by the Hawai‘i

Undersea Research Laboratory has been

jeopardized if not ended as a result of budget

cuts.

Beginning in 2015, NOAA’s Deep Sea Coral

Research and Technology Program, in

partnership with NOAA’s Office of Ocean

Exploration and Research, National Marine

Sanctuary Program, and NMFS, began a major

three‐year field research initiative, CAPSTONE,

in the Pacific Islands region. This campaign

conducted the first major biogeographic

exploratory work on deep‐sea coral and sponge

ecosystems outside the Hawaiian Archipelago,

as well as ecological studies off Hawai’i , where

the infrastructure and technical capacity exist to

support this type of field work. The results will

provide information on monument resources

and increase scientific understanding of these

unique deep‐sea ecosystems.

Online Annex – Comprehensive list of deep‐

sea corals in the U.S. Pacific Islands region

linked here:

https://deepseacoraldata.noaa.gov/library/2017‐

state‐of‐deep‐sea‐corals‐report

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Baco AR, Shank TM (2005) Population genetic

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Davies AJ, Guinotte JM (2011) Global habitat

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Figueroa D, Baco AR (2014) Complete mitochondrial

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Figueroa D, Baco AR (2015) Octocoral mitochondrial

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reversals, and also into the use of mitochondrial

genomes for cnidarian phylogenetics. Genome

Biology and Evolution 7(1):391‐409

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

Gazave E, Lapebie P, Renard E, Vacelet J, Rocher C,

Ereskovsky AV, Lavrov DV, Borchiellini C (2010)

Molecular phylogeny restores the supra‐generic

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Houlbrèque F, McCulloch M, Roark B, Guilderson T,

Meibom A, Kimball J, Mortimer G, Cuif J‐P,

Dunbar R (2010) Uranium‐series dating and

growth characteristics of the deep‐sea

scleractinian coral: Enallopsammia rostrata from

the Equatorial Pacific. Geochimica et

Cosmochimica Acta 74:2380‐2395

Long DJ (2011) Distinct megabenthic structure‐

forming communities within the Makapuʹu deep‐

sea coral bed and shifts in crinoid abundance.

MS, Florida State University, 89pp

Long DJ, Baco AR (2014) Rapid change with depth

in megabenthic structure‐forming communities

of the Makapuʹu deep‐sea coral bed. Deep‐Sea

Res Pt II 99:158‐168

Morgan NB (2013) Community structure of benthic

invertebrates on Necker Ridge in the North

Pacific: Distribution, diversity, and preliminary

comparison to the Hawaiian Archipelago. M.S.

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Morgan NB, Cairns S, Reiswig H, Baco AR (2015)

Benthic megafaunal community structure of

cobalt‐rich manganese crusts on Necker Ridge,

North Pacific Ocean. Deep‐Sea Res. I. 104:92‐105

Morrison CL, Baco AR, Nizinski MS, Coykendall

DK, Demopoulos, AWJ, Cho W, Shank TM (this

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Opresko DM (2009) A new name for the Hawaiian

antipatharian coral formerly known as Antipathes

dichotoma (Cnidaria: Anthozoa: Antipatharia)1.

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Opresko DM, Wagner D, Montgomery AD, Brugler

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DeMello J, Breuer E (2014) NOAA Deep‐sea

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Prouty NG, Roark EB, Buster NA, Ross SW (2011)

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REGION

  

Prouty NG, Roark EB, Andrews AH, Robinson LF,

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Roark EB, Guilderson TP, Dunbar RB, Fallon SJ,

Mucciarone DA (2009) Extreme longevity in

proteinaceous deep‐sea corals. Proceedings of the

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Rogers AD, Baco AR, Griffiths H, Hart T, Hall‐

Spencer JM (2007) Corals on seamounts. In:

Pitcher TJ, Morato T, Hart PJB, Clark MR,

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Online Annex 1. Deep-sea Coral Taxa in the U.S. Pacific Islands Region: Depth and 

Geographic Distribution 

204
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  

Appendix – Distribution maps for the Hawaiian Archipelago and Johns

Map 1. Locations of black corals (Order Antipatharia) recorded in the National Deep‐Sea Coral and Sponge Database (as o

205
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REG

  

Map 2. Locations of branching stony corals (Order Scleractinia) recorded in the National Deep‐Sea Coral and Sponge Dat

2017).

206
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REG

  

Map 3. Locations of non‐gorgonian alcyonaceans (Order Alcyonacea) recorded in the National Deep‐Sea Coral and Spong

October 2017).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REG

  

Map 4. Locations of gorgonians (formerly Order Gorgonacea)) recorded in the National Deep‐Sea Coral and Sponge Datab

2017).

208
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REG

  

Map 5. Locations of sea pens (Order Pennatulacea) recorded in the National Deep‐Sea Coral and Sponge Database (as of O

209
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REG

  

Map 6. Locations of lace corals (Family Stylasteridae) recorded in the National Deep‐Sea Coral and Sponge Database (as of 

210
STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE U.S. PACIFIC ISLANDS REG

  

Map 7. Locations of sponges recorded in the National Deep‐Sea Coral and Sponge Database (as of October 2017). Note: Th

additional HURL records shown in Figure 6.

211
                       

                     

                       

                               

                           

Maxent predictive habitat model for the primary framework‐forming deep‐sea coral in the

Southeast U.S., Lophelia pertusa. Warmer colors indicate greater likelihood of suitable habitat,

determined by a cross‐validation method that determined likelihood thresholds using the ratio

of false positive to false negative classification results when the model was tested on data left

out of the fitting process (Kinlan et al. 2012a).

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •

212
213

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •

PREDICTIVE HABITAT 

MODELING FOR 

DEEP-SEA CORALS 

IN U.S. WATERS

I. Introduction

Predictive habitat modeling (PHM) is a cost-effective 

method for extending the range and utility of expensive 

and time consuming field efforts to identify deep-sea coral 

habitat in locations that have not been sampled. PHM 

is not intended and should not be used as a substitute 

for field surveys via remotely operated vehicles (ROVs), 

submersibles, and similar instruments. Instead, modeling 

is one component of a highly integrated process that 

includes biological surveys, oceanographic data gathering 

efforts, management and conservation actions, and new 

exploration and scientific efforts. New data collection and 

verification of model results generates feedback that allows 

for the creation of better models (as illustrated by the 

conceptual model in Figure 1). Recognition of the utility of 

regional-scale habitat suitability models for deep-sea corals 

is increasing both in domestic and international waters and 

the models have become accepted as a useful indicator of 

vulnerable marine ecosystems.

PHM complements field efforts by facilitating habitat 

predictions across large regions, and this method can 

be used to help focus limited resources in areas that 

have the highest probability of supporting deep-sea 

coral ecosystems. Predictive habitat models also provide 

important insights into the environmental conditions 

controlling deep-sea coral distribution (feedbacks in Figure 

1). Threats to deep-sea coral habitats such as climate 

change and the looming threat of ocean acidification 

reinforce the need to gain a better understanding of the 

physical, chemical, and oceanographic conditions that 

influence deep-sea coral survival. 

John M. Guinotte1, 

Sam Georgian2*, 

Brian P. Kinlan3, 

Matthew Poti3, 

and Andrew J. 

Davies4

• • •

1 Marine Conservation 

Institute (current affiliation: 

Ecological Services, U.S. 

Fish and Wildlife Service, 

Lakewood, CO)

2 Marine Conservation 

Institute, Seattle, WA

* Corresponding Author:

Samuel.Georgian@marineconservation.org

3 NOAA Marine Spatial 

Ecology Division, National 

Centers for Coastal Ocean 

Science, Silver Spring, MD

4 School of Ocean Sciences, 

Bangor University, U.K.
PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •

214

In this spotlight we provide an overview of 

regional-scale modeling efforts in U.S. waters, 

some of the methods used to determine 

predicted habitat for deep-sea corals, the utility 

of these efforts for researchers and resource 

managers, and the limitations of such models.

II. Model Methodology

Predictive habitat modeling (often called 

habitat suitability modeling, ecological niche 

modeling, or species distribution modeling) 

integrates the spatial distribution of coral 

colonies with environmental data (including 

geomorphology of the seafloor and physical, 

biological, and chemical variables) to estimate 

the potential niche and distribution of deepsea corals. 

Most deep-sea coral PHM studies to-date have 

used “presence-only” approaches, as the vast 

majority of data on coral distribution consists 

only of records of presence (occurrence). 

The range of environmental conditions for 

known coral locations is used to determine 

an “environmental envelope” which is then 

mapped across the entire region, including 

areas where no sampling has taken place. 

This process results in an index of relative 

habitat suitability, typically ranging from 0-1 

or 0-100, that predicts the relative likelihood 

that a given area harbors deep-sea coral 

habitat. Presence-only modeling approaches 

that have been commonly used have included 

maximum entropy modeling (Maxent; e.g., 

Davies and Guinotte 2011, Rengstorf et al. 

2012, Yesson et al. 2012, Rengstorf et al. 2013, 

Figure 1. Conceptual process diagram showing the inputs and outputs of predictive habitat models for 

deep-sea corals, including feedbacks from model outputs to exploration and scientific efforts that lead to 

continual improvement in model quality, as well as informed management, conservation, and spatial 

planning actions.
215

Georgian et al. 2014), Ecological-Niche Factor

Analysis (ENFA; eg., Leverette and Metaxas

2005, Bryan and Metaxas 2007, Davies et al.

2008, Dolan et al. 2008, Tittensor et al. 2009,

Taylor et al. 2013), and Genetic Algorithm for

Rule Set Production modeling (GARP; e.g.,

Guinan et al. 2009, Tong et al. 2013). Of these,

Maxent is becoming increasingly common,

as it has consistently outperformed other

presence-only techniques (Elith et al. 2006,

Tittensor et al. 2009, Tong et al. 2013).

Models that incorporate known absences

in addition to known presences are being

increasingly used to predict deep-sea

coral habitats. These “presence/absence”

approaches have included Boosted Regression

Trees (BRT; Tracey et al. 2011), Generalized

Linear Models (GLM; e.g., Woodby et al.

2009), Generalized Additive Models (GAM;

e.g., Ross and Howell 2013, Rooper et al.

2014), and logistic regression and Generalized

Estimating Equation models (GEE; e.g.,

Woodby et al. 2009). When both presences

and absences are recorded with reasonable

certainty, presence/absence models can

produce more accurate and therefore, more

useful habitat predictions. It is important

to note however, that absence data can be

misleading in models with coarse spatial

resolutions (> 200 m), as it cannot be assumed

that coral is absent from a large area unless

the entire area has been surveyed. Current

sampling methods in the deep-sea often

give us confirmation of coral absence in

only a small portion of each modeled cell. In

addition, absence data may be misleading if

the species’ distribution is not in equilibrium,

if dispersal is limiting, or due to historical

artifacts including population losses due to

human activities (Hirzel et al. 2002).

Difficulties in determining ‘true’ absences 

have led researchers to generate ensemble 

Recent trends in deep-sea 

coral modeling

1. Predictive habitat modeling for deepsea corals has increased in the last ten

years due to: a) improved resolution

and availability of environmental data,

b) increased quality and quantity of

coral data, c) increased recognition

that modeling is a useful and costeffective means to identify vulnerable

benthic habitats and d) increased

computational power and the

availability of appropriate algorithms

and software for predictive habitat

modeling.

2. For the first time, global and regional

modeling efforts can predict habitat

at scales that are biologically relevant

as well as practical for resource

management (10s of meters to 10s of

kilometers).

3. Presence-only modeling approaches

(e.g., Maxent, ENFA) have been

the most frequently used method

to model deep-sea coral habitat at

regional scales, but they do not show

probability of occurrence. Improved

sampling and analysis methods

are required to allow for presence/

absence models in the deep-sea. The

next generation of models should

incorporate measures of abundance

(e.g., biomass, number of colonies,

percent cover), and move beyond

presence-absence approaches.

4. Modeling has helped identify

important environmental correlates

of deep-sea coral distribution, which

is useful for forecasting areas where

corals are most at risk from climate

change and ocean acidification.

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •
216

models produced by using a combination 

of presence-only and presence/absence 

approaches (e.g., GAMs and Maxent; Ross 

and Howell 2013) to identify potential habitat. 

In the future, as databases improve, more 

data on deep-sea coral presence/absence 

and abundance (e.g., biomass, number of 

colonies, percent cover) will facilitate more 

sophisticated models (e.g., Guinan et al. 

2009). Models that can predict areas of high 

abundance or biodiversity hotspots are of 

particular importance to conservation and 

resource managers, who must weigh the 

biological and ecological value of different 

areas against economic costs associated with 

their protection (Ardron et al. 2014). 

Field survey efforts offer critical support to 

these improved datasets and help to avoid 

one of the primary pitfalls of presence-only 

models: important, undiscovered coral 

habitats can easily be missed because the 

presence locations are unlikely to fully 

represent the range of possible deep coral 

habitats. Verified absence data should 

always be preferred over a model prediction 

of “unsuitable habitat.” The collection of 

georeferenced, verified absence data with 

broad spatial coverage would greatly reduce 

sampling bias and help to improve the 

next generation of models. Yet, due to the 

time and expense involved, deep-sea field 

surveys are rarely randomly distributed 

spatially, introducing sampling bias that will 

always be an important consideration in the 

interpretation of model results. The adage by 

Carl Sagan that “absence of evidence is not 

the evidence of absence” is especially critical 

in interpretation of deep-sea field survey data 

and model outputs (discussed in Etnoyer 

and Morgan 2007). Habitat suitability models 

can influence spatial management decisions, 

so they should be tested for their validity, 

updated, and improved periodically.

Models also differ in the types of

environmental data they use to make

predictions. At a minimum, environmental

data usually include bathymetry (depth)

and statistics derived from bathymetry that

characterize the topography of the seafloor,

generally referred to as terrain metrics.

Other types of environmental variables

often included in models are substrate

(benthic sediment types and the distribution

of hard bottom) and oceanographic data.

Oceanographic data encompasses physical

variables (e.g., temperature, salinity, and

currents), biological variables (e.g., surface

productivity and particulate organic carbon

flux to the seafloor), and chemical variables

(e.g., pH, dissolved oxygen, and carbonate

chemistry). The importance of individual

environmental variables in determining the

distribution of deep-sea corals appears to vary

considerably both among taxa and among

regions (Table 1).

Seven environmental variables that are

consistently strong predictors across

regions and taxa include: depth, seafloor

geomorphology (slope, curvature, roughness,

changes in slope), sediment/substrate type,

carbonate chemistry, temperature, salinity,

indices of near-bottom current velocity,

and food flux to benthic environments. The

regional variation in approach and data

priority in modeling efforts often depends

in part on the types of data are available.

For example, carbonate chemistry has been

found to be among the most important

predictor variables for the global occurrence

of both scleractinian reef-forming corals

and octocorals (Davies and Guinotte 2011,

Yesson et al. 2012), but deep-water carbonate

chemistry data are limited or non-existent in

many regions. Other variables are proxies for

the actual processes driving coral distribution;

for example, near-bottom currents are thought

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •
Study Model Taxa Resolution Terrain Substrate Chemical 

Northeast and Mid-Atlantic U.S.Kinlan et al. in press Maxent Alcyonacea, Gorgonacea, Pennatulacea, Sessiliflorae, Scleractinia, 

Caryophylliidae, Flabellidae

370 m Aspect, depth, SL, SL‐of‐SL, 

rugosity,

PLC , PRC, BPI

Mean grain size, % sand, 

%mud, %gravel

– 

s

Bryan and Metaxas 2007

ENFA Paragorgiidae, Primnoidae 5 km Depth, SL – – C

Southeast U.S.Davies (unpublished), Mienis et al. 2014

Maxent Lophelia pertusa, Madrepora oculata, 

Enallopsammia profunda, Solenosmillia variabilis, Oculina varicosa

90 m, 1 

km

Depth, SL, rugosity SL-ofSL, PLC, PRC, BPI – ΩA, nitrates 

Kinlan et al. 2012a Maxent Lophelia pertusa, Madrepora oculata, 

Enallopsammia profunda, FF Scleractinia (exc. Oculina spp.), Non-FF Scleractinia, Antipatharia, Stylasteridae, Pennatulacea, Alcyonacea, Gorgonacea, Holaxonia, Calcaxonia

370 m Depth, SL, SL-of-SL, aspect, 

rugosity, PLC, PRC, BPI

Mean grain size, % sand, 

%mud, %gravel

– 

s

Gulf of MexicoGuinotte and Davies (in prep)

Maxent Lophelia pertusa, Madrepora oculata, 

Enallopsammia profunda, Solenosmillia variabilis, Oculina varicosa

~90 m Depth, SL, rugosity To be determined ΩA, ΩC 

sa

Georgian et al. 2014 Maxent Lophelia pertusa 5 m, 25 m Depth, SL, roughness, TPI, 

LonC, LatC, GC, eastness, 

northness

Potential hard-bottom locations from 3D seismic data ΩA 

Kinlan et al. 2012b Maxent Lophelia pertusa, Madracis, Madrepora, Bebryce, Callogorgia, Hypnogorgia, 

Paramuriceidae, Plexauridae, Isididae, Ellisellidae, Chrysogorgiidae, 

Antipatharia, Alcyonacea, Gorgonacea, Scleractinia, FF Scleractinia, Non-FF Scleractinia, All FF taxa

370 m Depth, SL, SL-of-SL, aspect, 

rugosity, PLC, PRC, BPI

Mean grain size, % sand, 

% mud, % gravel, potential hard-bottom locations 

interpreted from 3D 

seismic data

– 

s

West Coast U.S.Guinotte and Davies 2012, 2014

Maxent Antipatharia, Scleractinia, Alcyoniina, Calcaxonia, Holaxonia, 

Scleraxonia

500 m Depth, SL – ΩA, ΩC, 

nitrate, 

phosphate, 

silicate

Huff et al. 2013 GAM, 

GLM

Antipathes dendrochristos 90 m Depth, SL, PRC – – T

lin

v

Table 1. Summary of key features of the regional and global-scale deep-sea coral predictive habitat models discussed in this chapter, incluthe modeling approach used, the spatial resolution, and the environmental variables used as predictors. Environmental variables that werare italicized. Abbreviations: ENFA=Environmental Niche Factor Analysis, Maxent=Maximum Entropy, GAM=Generalized Additive Mabbreviations: Chl. a=chlorophyll a, CPI=Chlorophyll Persistence Index, BPI=bathymetric position index, POC=particulate organic carbosaturation state, ΩC=calcite saturation state, TPI=topographic position index, AOU=apparent oxygen utilization, TA=total alkalinity, [Cconcentration, DIC=dissolved inorganic carbon, lat./long.=latitude/longitude, TRI=terrain ruggedness index, DO=dissolved oxygen, SL=slope, PLC=plan curvature, PRC=profile curvature, LonC=longitudinal curvature, LatC=latitudinal curvature, GC=general curvature, FVariables are at-depth unless otherwise indicated (i.e., surface chlorophyll a, surface primary production, surface turbidity).
Bryan and Metaxas 

2007

ENFA Paragorgiidae, Primnoidae 5 km Depth, SL – – C

AlaskaGuinotte and Davies 2013

Maxent Alcyoniina, Antipatharia, Calcaxonia, Filifera, Holaxonia, Scleractinia, Scleraxonia, Stolonifera 700 m Depth, SL, rugosity, BPI, PLC, PRC, tangential curvature, aspect, northness, eastness, TRI, TPI, 

roughness

– ΩA, ΩC, 

nitrate, 

phosphate, 

silicate

Bryan and Metaxas 2007

ENFA Paragorgiidae, Primnoidae 5 km Depth, SL – – C

Rooper et al. 2016 GAM Primnoidae, Stylasteridae, all corals 

combined, Porifera 

100 m 

(Aleutian 

Islands), 

1km (Bering Sea)

Depth, SL, rugosity, lat./long., 

aspect 

Sediment type – 

cu

Sigler et al. 2015 GAM Plexauridae,Primnoidae, Isididae, 

Paragorgiidae, Antipatharia, Pennatulacea, Porifera

1 km Depth, SL, lat./long. Grain size, sediment sorting – 

Hawaii and U.S. Pacific TerritoriesBauer et al. 2016 Maxent Calcaxonia, Holaxonia, Scleraxonia, 

gold corals, red and pink corals, black corals, bamboo corals,

bubblegum corals, FF and non‐FF stony corals, gorgonian and non‐gorgonian soft corals

360 m Depth , SL, SL‐of‐SL , 

aspect, rugosity, total 

curvature , PLC, PRC, BPI, 

distance to shore , distance 

to seamounts

– – 

s

c

cu

GlobalDavies and Guinotte 2011

Maxent FF deep-sea corals (Enallopsammia 

rotrata, Goniocorella dumosa, Lophelia pertusa, Madrepora oculata, Solenosmillia variabilis)

~1 km Depth, SL, rugosity – TA, pH, 

[CO32-], 

ΩA, ΩC, 

nitrate, 

phosphate, 

silicate, 

DIC

R

D

Yesson et al. 2015 Maxent Antipatharia (Antipathidae, 

Aphanipathidae, Cladopathidae, Leiopathidae, Myriopathidae, Schizopathidae, Stylopathidae)

1 km Curvature, SL – TA 

Yesson et al. 2012 Maxent Alcyoniina, Calcaxonia, Holaxonia, 

Scleraxonia, Sessiliflorae, Stolonifera, Subselliflorae

~1 km Depth, SL, rugosity – TA, pH, 

[CO32-], 

ΩA, ΩC, 

nitrate, 

phosphate, 

silicate, 

DIC

Tittensor et al. 2009 Maxent, 

ENFA

Scleractinia corals on seamounts ~130 km Depth – TA, ΩA, 

nitrate, 

phosphate, 

silicate, 

total DIC

r

Davies et al. 2008 ENFA Lophelia pertusa 0.25°, 1° Depth, aspect, SL Hydrocarbon seeps 

and pockmarks, iceberg 

ploughmarks

TA, ΩA, 

DIC, 

nitrate, 

phosphate, 

silicate

C
219

to be very important determinants of particle 

flux, coral distribution and abundance 

(White et al. 2007, Mienis et al. 2012), but 

data on bottom currents are limited. Terrain 

metrics may therefore serve as proxies 

for currents. Moreover, the importance of 

geomorphological variables such as slope 

and curvature appears to be highly scaledependent, with high-resolution multibeam 

bathymetry often required to reveal fine-scale 

seafloor features associated with suitable 

coral habitat. More in situ observation 

and experimentation, more consistent 

collection and integration of high-resolution 

bathymetric and seafloor characterization 

products, and better data-assimilating deep 

ocean biophysical and chemical models are 

necessary to better resolve questions about the 

primary drivers of deep-sea coral distribution 

and abundance. 

Finally, models vary in the level of spatial and 

taxonomic detail, or resolution, they convey. 

The spatial resolution of predictive habitat 

modeling is often limited by the availability 

of fine-grained spatial data for environmental 

variables. For example, the spatial scales of 

environmental data may be coarse (1,000-

10,000 meters resolution) compared to 

the resolution of multibeam bathymetry 

available in the same study area (2-40 m 

resolution, depending on depth). This issue, 

combined with the reality that high-resolution 

multibeam does not exist across entire study 

areas, often leads modelers to predict habitat 

at coarser resolutions than the multibeam 

bathymetry. Regional fisheries management 

councils, conservation organizations, and 

habitat suitability modelers seek to manage 

areas larger than the extent of fine-scale data 

that are available, so such a compromise is 

necessary. There is a need for more extensive

spatial coverage of fine-scale resolution

environmental data, and conversely, a need

for more habitat suitability studies that

operate over smaller extents. Local-scale

studies with fine-resolution data allow

researchers to incorporate locally important

drivers of species distributions, like bottom

hardness and local relief. One large boulder,

or one meter of vertical relief can make a

considerable difference in habitat quality for

deep-sea corals and sponges.

Difficulties with species identification can

inhibit taxonomic resolution: it is not always

possible to model habitat for a single coral

species because the observations are recorded

at the genus or family level. In many cases,

useful models are created by grouping

coral species at higher taxonomic levels,

ranging from genus to family, suborder, or

order. Grouping to high taxonomic levels

is often necessary from a practical resource

management perspective when managers

cannot deal with hundreds of species

models. However, careful interpretation and

assessment of predictions should be exercised

when members of those taxa occupy very

different environmental niches (e.g., Quattrini

et al. 2013). In other cases, corals may be

grouped for modeling by their functional

similarity, for example, branching stony corals

that form a rigid framework and thus form

habitat for fishes and invertebrates (e.g., Dolan

et al. 2008).

Below, we review recent and ongoing

predictive habitat modeling efforts in different

regions of the U.S.,1

 providing a brief overview

of approaches, data, taxa modeled, resolution,

results, and management/conservation

applications in each region.

1

There are many fine-scale PHM efforts underway or completed for deep-sea coral habitat for 

specific sites in U.S. waters. These efforts are not reviewed here.

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •
220

III. Predictive Habitat

Modeling in U.S. Regions

III.1. Northeast and Mid-Atlantic red

Over the past 5 years, NOAA’s Deep-sea 

Coral Research and Technology Program 

has supported development of a geospatial 

database of known coral presence locations 

in the Northeast and Mid-Atlantic regions 

(e.g., Packer et al. 2007, Scanlon et al. 2010, 

Packer and Dorfman 2012, Packer and Drohan 

2012, NOAA 2015). The New England Fishery 

Management Council has used these locations, 

in conjunction with primary literature, expert 

opinion, and geomorphological characteristics 

of canyons, to help identify and prioritize 

known and potential areas of deep-sea coral 

habitat (NEFMC 2012). This constitutes a first 

step toward predictive modeling and has 

helped the Council as they consider various 

alternatives for coral habitat protection. 

NOAA scientists also produced a suite of 

moderate resolution (~370 m) models for 

multiple deep-sea coral taxonomic groupings 

in the northeast and Mid-Atlantic region 

(Kinlan et al. in prep; Figures 2 and 3). The 

inclusion of multiple spatial scales was 

designed to capture biologically-relevant 

features ranging from relatively fine-scale 

topography, the presence of deep-water 

canyons, and the continental shelf break. 

Maxent models were constructed using 

presence-only data and environmental 

variables including depth, terrain metrics 

calculated at multiple spatial scales (slope, 

slope-of-slope, rugosity, plan curvature, 

profile curvature, and bathymetric position 

index), substrate (mean grain size and percent 

sand/mud/gravel of benthic sediments), 

and physical (bottom temperature, salinity, 

and surface turbidity), biological (surface 

chlorophyll-a), and chemical (dissolved 

oxygen) oceanographic variables. 

In order to improve model parsimony while

maintaining predictive power, a stepwise

model selection process was employed to

uce the final number of environmental

variables included in the models by removing

the most redundant variable remaining at each

step. The model runs from the model selection

process were ranked from best to worst by

model performance and by model complexity,

and the model run with the best average rank

was selected as the final model. To generate

model predictions in a format that can be

directly compared across taxonomic groups

and regions, Maxent outputs were classified

into habitat suitability likelihood classes using

breakpoints corresponding to ratios of the cost

for false positive errors versus the cost for false

negative errors.

Three major taxonomic groups were modeled:

stony corals (order Scleractinia), sea pens

(order Pennatulacea), and other octocorals

(orders Alcyonacea and Gorgonacea

combined). For each major taxonomic group,

two subgroups were also considered based

on suborder or family-level taxonomy. The

Gorgonacea subgroup incorporates most of

the potential habitat-forming species in this

region (Packer et al., this volume).

The models predicted extensive areas of

potential habitat on the continental slope for

all major taxonomic groups and subgroups

modeled. Highest suitability values were

concentrated in submarine canyon areas at

depths of 300-2000 m. Results were used to

identify several unexplored potential hotspots

of deep coral habitat suitability in the MidAtlantic and New England canyons, and

subsequent ground-truthing cruises from

2012-2015 aboard NOAA Ship Henry Bigelow

and NOAA Ship Okeanos Explorer (Nizinski

and Shank 2012, NOAA OER, 2013, 2014,

NRDC 2014, Quattrini et al. 2015) confirmed

the existence of these hotspots (e.g., Figure

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • • • •
221

3). Efforts are underway to improve the 

resolution and accuracy of this model using 

coral data collected during these field efforts 

and multibeam bathymetry collected as part 

of the Atlantic Canyons Undersea Mapping 

Expeditions (ACUMEN) project (NOAA OER 

2012), in a direct application of the process 

model illustrated in Figure 1. 

Predictive habitat modeling helped guide 

BOEM-funded field surveys of canyons in the 

Mid-Atlantic region in 2012 (Ross and Brooke 

2012). Using archived historical observations 

from surveys in the 1980s in conjunction with 

modern state-of-the-art multibeam, areas 

of potential hard ground and coral habitat 

were modeled from a suite of terrain and 

environmental variables within Mid-Atlantic 

canyons using the Maxent approach. This 

approach highlighted the value of local-scale, 

high-resolution models in guiding and focusing 

Figure 2. Environmental layers used in the creation of Maxent deep-sea coral predictive habitat models for the 

Northeast U.S. region. A) bathymetric depth, B) bathymetric position index (BPI) calculated at a 20 km scale, C) 

predicted annual bottom temperature, D) predicted surficial sediment mean grain size, E) slope of slope, F) surface 

turbidity. 

A B C

D E F

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •
222

Figure 3. Maxent predictive habitat model for Gorgonian Alcyonaceans in the Oceanographer, Gilbert, 

and Lydonia Canyon Complex in the Northeast U.S. region. Black crosses indicate locations of known 

coral locations discovered during a 2012 ground-truthing cruise aboard the NOAA Ship Henry

Bigelow (Nizinski and Shank 2012). FPR=False positive rate. FNR=false negative rate.

research effort within a clearly defined area 

whilst utilizing past observations. The models 

have been validated in the field and visual 

observations collected during subsequent 

cruises will be used to enhance the local-scale 

model. 

All of the models described will continue to 

be developed and refined, however they have 

already played a seminal role in regional 

fishery management. In 2015, the Mid-Atlantic 

Fishery Management Council (MAFMC 2015) 

recommended establishment of “deep-sea 

coral zones” to protect deep-sea corals from 

the impacts of bottom-tending fishing gear. 

This fishery management plan amendment 

was recently approved by the National Marine 

Fisheries Service, and will protect over 98,000 

km2

 of habitat in canyons and deep-water 

areas where corals have been observed or 

where they were predicted to occur based 

on NOAA Maxent models. This was among 

the first explicit examples of deep-sea coral 

predictive habitat models playing a major 

role in U.S. fishery management conservation 

decisions, confirming the utility of PHM for 

future conservation and management efforts. 

The New England Fishery Management 

Council is expected to use similar approaches 

in a planned deep-sea coral amendment. 

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •
223

III.2. Southeast

In 2009, Davies (unpublished) created the 

first Maxent predictive habitat model in the 

Southeast U.S. Atlantic region for frameworkforming scleractinian corals (including 

records for Lophelia pertusa, Madrepora oculata, 

Enallopsammia profunda, Solenosmilia variabilis

and Oculina varicosa). Environmental variables 

were created following procedures later 

described in Davies and Guinotte (2011) using 

the highest resolution bathymetry available 

including the National Geophysical Data 

Center’s Coastal Relief Model at 3 arc second 

(~90 meter) resolution and global bathymetry 

available at 30 arc second (~1 km) resolution 

(Becker et al. 2009). Relevant environmental 

layers were selected for the model, including 

omega aragonite (Orr et al. 2005), depth 

(Becker et al. 2009), dissolved oxygen (Garcia 

et al. 2006), salinity (Boyer et al. 2009), 

nitrate concentrations (Garcia et al. 2006), 

temperature (Boyer et al. 2009) and rugosity 

(Wilson et al. 2007). Model results from this 

effort were overlaid with existing fishery 

closures to identify areas with high habitat 

suitability that remain at risk from destructive 

fishing practices.

In 2012, NOAA produced a set of 370 meter 

resolution regional predictive habitat models 

for deep-sea coral taxonomic groups in the 

Figure 4. Maxent predictive 

habitat model for all frameworkforming Scleractinian deep-sea 

corals in a subset of the U.S. 

Southeast Atlantic. Warmer 

colors indicate greater likelihood 

of suitable habitat, determined 

by a cross-validation method that 

determined likelihood thresholds 

using the ratio of false positive to 

false negative classification results 

when the model was tested on data 

left out of the fitting process (see 

Kinlan et al. 2012b and David et 

al., this volume for details).

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •
224

Southeast U.S. (Kinlan et al. 2012a). Example 

outputs from this model are shown in Figure 

4 and in Hourigan et al. (this volume). 

Using a similar approach to that employed 

in the Northeast region, Maxent models 

were constructed using presence-only data 

and environmental variables selected from 

a set of candidate predictors. A total of 14 

taxonomic groups were modeled, including 

three important species of framework-forming 

stony corals (Lophelia pertusa, Madrepora 

oculata, and Enallopsammia profunda), an 

important framework-forming genus (Oculina

spp.), all framework-forming stony corals as 

a group (Figure 4), non-framework-forming 

stony corals, black corals, lace corals, sea pens, 

gorgonian and non-gorgonian soft corals, and 

two suborders of gorgonian corals (Holaxonia, 

Calcaxonia). The models predicted extensive 

areas of potential framework-forming deep 

coral habitat, concentrated at depths of 100-

1000 m. Results confirmed that existing coral 

protection areas covered the majority of 

likely framework-forming deep coral habitat 

(Figure 4). The models were provided to the 

South Atlantic Fishery Management Council 

to support discussions of Ecosystem Based 

Management measures including a recent 

amendment to the Fishery Management Plan 

for Coral, Coral Reef, and Live/Hardbottom 

Habitats in the South Atlantic Region (SAFMC 

2013). 

The two Southeast models highlight the issue 

that model results can vary considerably in 

both the extent of highly suitable habitat and 

in the suitability scores of individual grid 

cells, even when similar modeling methods 

are used (i.e., Maxent). Predictor variables 

used as input in the models are likely 

responsible for these differences (e.g., seafloor 

substrate type), but the coral records used 

to determine the environmental niche also 

play an important role in determining final 

model results. Multiple independent modeling 

efforts are useful because they can highlight

areas of high certainty (where multiple

models agree), reveal sensitivity of model

predictions to different assumptions, methods,

and input data, and improve future model

iterations. Determination of which model(s)

more accurately predicts reality can only be

accomplished through field validation efforts.

III.3. Gulf of Mexico

Marine Conservation Institute and Bangor 

University are currently developing Maxent 

models for both deep-sea and mesophotic 

coral habitat in the Gulf of Mexico. The spatial 

resolution of model results is based on the 

National Geophysical Data Center’s Coastal 

Relief Model at 3 arc second (~90 meter) 

resolution. Modeling methods are similar to 

the approach used in the Southeast U.S, U.S. 

West Coast, and Alaska. The objectives of 

this work are to identify both deep-sea and 

mesophotic reef habitat that are not currently 

under protection from human activity 

including oil and gas production/accidents, 

bottom trawling, and climate change. 

The Gulf of Mexico presents unique challenges 

and opportunities for predictive coral habitat 

modeling. High-resolution seismic data that 

can be used to detect potential hard-bottom 

patches are lacking in most of the U.S. 

Exclusive Economic Zone (EEZ), but this is not 

the case in the Gulf of Mexico due to a long 

history of oil and gas exploration in the region. 

Extensive 3D seismic surveys conducted over 

several decades in large areas of the Gulf 

have recently been interpreted to provide 

high-resolution information on potential 

hardground areas that are helping to improve 

predictions of coral habitat suitability (Boland 

et al., this volume). 

In 2012, scientists at NOAA applied the 

Maxent modeling approach used in the 

Northeast and Southeast regions to develop 

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •
225

moderate spatial resolution (370 meter) 

regional models for deep-sea coral taxonomic 

groups in the northern Gulf of Mexico (Kinlan 

et al. 2012b). Maxent models were constructed 

using presence-only data and environmental 

variables selected from a suite of terrain, 

substrate (including potential hardground 

locations), and physical and biological 

oceanographic variables. Taxonomic groups 

similar to those used in the Southeast region 

were used for modeling, with an emphasis 

on predictions of key framework-forming 

species. This work has already been used in 

targeting exploration surveys and is expected 

to contribute to efforts at habitat protection, 

spatial planning, and fishery management in 

the Gulf of Mexico. 

In 2014, Georgian et al. developed highresolution (25 m) Maxent models predicting 

the distribution of Lophelia pertusa across a 

large region of the northern Gulf of Mexico. 

The authors used a suite of 11 environmental 

variables including depth, a number of 

terrain metrics, omega aragonite, substrate 

type, and export productivity. Substrate 

type consisted of potential hardground areas 

developed by the Bureau of Ocean Energy 

Management (Boland et al., this volume) and 

was highly predictive of L. pertusa distribution, 

demonstrating the utility of these type of 

data for future PHM efforts. The model 

was tested during a field survey aboard EV 

Nautilus by Ocean Exploration Trust in 2013 

and successfully predicted the location of two 

large L. pertusa mounds. 

III.4. U.S. Caribbean

There have been no predictive habitat models 

for deep-sea corals in the U.S. Caribbean 

(Puerto Rico, U.S. Virgin Islands, and Navassa 

Island), despite their relatively high diversity 

in the region (Cairns 1979, 2007). While several 

global-scale modeling efforts have included 

parts of the Caribbean (e.g., Tittensor et al. 

2009, Davies and Guinotte 2011), there is 

an urgent need to better characterize the 

distribution of cold-water corals throughout 

the region. 

III.5. West Coast (Washington,

Oregon, California)

Bryan and Metaxas (2007) published the first 

predictive habitat models for deep-sea corals 

(Families Paragorgiidae and Primnoidae) 

for the West Coast and Alaska. Four years 

later, in 2011, predictive habitat models 

were developed for six taxonomic groups 

of deep-sea corals (Orders Antipatharia 

and Scleractinia, Suborders Alcyoniina, 

Calcaxonia, Holaxonia and Scleraxonia) 

in the waters of Washington, Oregon and 

California (Guinotte and Davies 2012, 2014). 

The objectives of this effort were to: 1) aid 

future research and mapping efforts for 

deep-sea coral habitats, 2) Assess potential 

coral habitat suitability both within and 

outside existing bottom trawl closures (e.g., 

to inform designation and protection of 

essential fish habitat – EFH), and 3) identify 

suitable habitat in and around the region’s 

six National Marine Sanctuaries. Maxent was 

used to model deep-sea coral habitat at a 

500 m spatial resolution using coral records 

collected from a variety of sources and a 

regional database of 30 physical, chemical 

and environmental variables. Figure 5 shows 

the model results for Washington waters with 

overlays of existing fishing closures and the 

Olympic Coast National Marine Sanctuary. 

These modeling results will be used to assess 

gaps in protection afforded by existing fishery 

management closures through the Pacific 

Fishery Management Council’s periodic 

review of essential fish habitat (Huff et al. 

2013; Guinotte and Davies, 2014). 

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III.6. Alaska

Several regional-scale predictive habitatmodeling efforts have been conducted or are 

currently underway for deep-sea coral and 

sponge habitats in Alaskan waters. Guinotte 

and Davies (2013) developed Maxent models 

for the Alaska EEZ at a spatial resolution of 

~700 m using 30 environmental, physical, 

and chemical predictor variables. Models at 

the Suborder and Order levels revealed that 

the majority of predicted suitable habitat 

for gorgonians and lace corals (Family 

Stylasteridae) occurs in the Aleutian Islands, 

and to a lesser extent the eastern Bering 

Slope, Gulf of Alaska seamounts, and the 

Fjord region and shelf break of Southeast 

Alaska. Soft corals (Suborders Alcyoniina 

and Stolonifera) additionally had predicted 

suitable habitat on the eastern Bering Sea 

Shelf. Guinotte and Davies (2013) also 

modeled a complex habitat type, coral and 

sponge “gardens” (Stone and Shotwell 2007) 

in the Aleutian Islands. While some coral 

and sponge gardens are currently protected 

from bottom-contact fishing, other areas of 

predicted suitable habitat remain open to such 

fishing. Results from this modeling effort will 

be combined with new information on the 

skeletal mineralogy of Alaska’s deep-sea corals 

(Stone et al. in prep.) and predicted changes in 

carbonate chemistry through 2100 to produce 

a spatially explicit ocean acidification risk 

assessment for Alaska’s coral resources.

NOAA’s Alaska Fisheries Science Center has 

also developed generalized additive models 

(GAMs) for coral and sponge habitat in Alaska 

Figure 5. Maxent predictive habitat 

model for all taxa (Orders: Antipatharia 

and Scleractinia; Suborders: 

Alcyoniina, Calcaxonia, Holaxonia, 

and Scleraxonia) for the Olympic Coast 

National Marine Sanctuary. Warmer 

colors indicate greater likelihood of 

suitable habitat across all taxa modeled 

(see Guinotte and Davies 2012 for 

details on likelihood). Hatched areas are 

existing trawl closures (EFH).

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

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227

that use data from bottom trawl surveys. 

Rooper et al. (2014) presented models that 

predicted presence or absence, abundance, 

and family diversity of corals and sponges 

(as opposed to presence-only models) for 

the Aleutian Islands. The predictor variables 

included remotely-sensed data, predictions 

from oceanographic models, and locationspecific data collected during trawl surveys. 

The PHM objective was to develop and 

parameterize a spatially explicit model to 

predict coral and sponge presence, abundance, 

and diversity at 100 m spatial resolution for 

the Aleutian Islands. Field validation surveys 

using towed camera systems and a stratifiedrandom station selection were conducted in 

2012-2014 and data are in the process of being 

analyzed. Similar models based on bottom 

trawl survey data have also been produced 

for corals and sponges in the Gulf of Alaska 

(Rooper et al. 2017). These models will be 

ground truthed using available underwater 

image data.

Using catches from trawl surveys and similar 

GAM techniques, Sigler et al. (2015) developed 

models of coral (excluding Pennatulacea), sea 

whip (Pennatulacea), and sponge distribution 

for the outer shelf and slope of the eastern 

Bering Sea. In 2014, randomized camera 

surveys were conducted to verify these 

distribution models (Rooper et al. 2016). Coral 

densities were low, but the model based on 

trawl survey data was generally reliable for 

predicting coral presence or absence in the 

camera survey. The bottom trawl survey 

models also successfully predicted sponge and 

sea whip presence or absence, but to a lesser 

degree than for coral. Presence or absence 

models of corals, sponges and sea whips were 

also constructed from the camera survey data. 

Combining these models with the distribution 

models constructed from bottom trawl survey 

data predicted the distribution of corals, 

sponges, and sea whips with better accuracy

than the individual models did during crossvalidation. These models and associated data

are being used by the North Pacific Fishery

Management Council in deliberations on

potential additional habitat protections in the

Eastern Bering Sea.

III.7. Hawaii and U.S. Pacific

Territories

In 2015, scientists at NOAA National Centers 

for Coastal Ocean Science applied the same 

Maxent modeling approach used in the 

Northeast, Southeast, and Gulf of Mexico 

to develop moderate resolution (360 meter) 

regional models for deep-sea coral taxonomic 

groups in the Main Hawaiian Islands (Bauer 

et al. 2016). Maxent models were constructed 

using presence-only data and a suite of 

environmental variables including depth, 

terrain metrics calculated at multiple spatial 

scales, proximity to seamounts, and physical 

and biological oceanography. A total of 16 

taxonomic groups were modeled, including 

gold corals, red and pink corals, black corals 

(separated into distinct groups by depth), 

bamboo corals, bubblegum corals, frameworkforming and non-framework-forming stony 

corals, gorgonian and non-gorgonian soft 

corals, three suborders of gorgonian corals 

(Calcaxonia, Holaxonia, Scleraxonia), and 

sea pens (separated into distinct groups 

by substrate). In addition, models were 

constructed to predict high diversity areas 

likely to support ≥ 4 and ≥ 7 genera of 

deep-sea corals. For many of the taxonomic 

groups, predictions of potential habitat were 

driven by depth. These models were part 

of a comprehensive marine biogeographic 

assessment to provide data products to inform 

the Bureau of Ocean Energy Management’s 

renewable energy policy decisions in Hawaii.

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228

While there are many coral occurrence records 

in the Hawaiian archipelago, information from 

other U.S. Pacific Territories is sparse (Parrish 

et al., this volume). The only modeling 

results for these areas are from globalscale efforts for 6 species of scleractinian 

reef formers (Davies and Guinotte 2011), 

octocorals (suborders Alcyoniina, Calcaxonia, 

Holaxonia, Scleraxonia, Sessiliflorae, 

Stolonifera, and Subselliflorae (Yesson et al. 

2012)), and black corals (Yesson et al. 2017). 

Both efforts modeled deep-sea coral habitat 

at a ~1 km spatial resolution using Maxent, 

global databases for coral locations, and 

30 environmental, physical, and chemical 

predictor variables.

IV. Application of Predictive

Habitat Models to Fishery

Management

In addition to helping researchers identify 

areas where coral habitat is most likely to 

be found, modeling results can be used 

in conjunction with existing fisheries 

management boundaries to help resource 

managers identify areas where potential 

deep-sea coral habitat remains at risk from 

human activity. Figure 5 shows the predicted 

distribution of several taxa of deep-sea corals 

and existing areas closed to bottom trawls to 

protect essential fish habitat for groundfish off 

Washington (Guinotte and Davies 2012, 2014). 

Commercial fishing using bottom-contact gear 

is a common practice in most U.S. regions, but 

field surveys for deep-sea corals are limited 

by the expense and extent of area that can 

be surveyed. Predictive habitat modeling 

represents a potential cost-effective means to 

fill these gaps. 

Modeling results can also be overlaid with 

coral bycatch maps to test the accuracy 

of model results. However, this type of

comparison can give conflicting results when

commercial fishers actively avoid high-relief

areas (most likely to harbor most deep-sea

corals) in order to minimize gear damage

and/or loss. Modeling results can and should

be incorporated into U.S. regional Fishery

Management Councils’ fisheries review

processes to help determine areas where

additional management measures may be

warranted or where existing boundaries need

to be amended.

For example, regional PHM results in

the Southeast U.S. Atlantic contributed to

discussions of alternatives in the development

of the SAFMC’s Coral Amendment 8 to the

Fishery Management Plan for Coral, Coral

Reef, and Live/Hardbottom Habitats in the

South Atlantic Region. This outcome resulted

in both the expansion of some Habitat Area

of Particular Concern (HAPC) boundaries

(Figure 4) to capture more high-likelihood

coral habitat, and opening of some flat bottom

areas of low likelihood to support coral habitat

(SAFMC 2013). In 2015, the Mid-Atlantic

Fishery Management Council approved the

closure of 15 submarine canyon and slope

areas to almost all bottom-contact fishing gear

specifically to protect deep-sea coral habitats

(MAFMC 2015). The results of the NOAA

deep-sea coral PHM were included in the

closure proposal and specifically used in the

identification of boundaries for these discrete

areas. Decisions like these are difficult to make

based on spatially limited field sampling

efforts alone; comprehensive regional maps

of habitat suitability produced by PHM can

greatly assist in developing and prioritizing

spatial boundaries. Similar discussions are

underway to varying degrees in other Fishery

Management Councils, including the New

England, Pacific and North Pacific Councils,

and PHM results are likely to be a useful tool

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229

for shaping and evaluating alternative spatial 

management and conservation measures in 

these regions.

V. Limitations of Predictive

Habitat Modeling Results

There are several limitations to predictive 

habitat modeling. Many of the variables that 

are important for coral settlement, growth 

and survival cannot be incorporated into the 

models because data on these variables do not 

exist at sufficient resolutions or are limited 

in geographic extent. This is particularly true 

for data on high-resolution current direction/

velocity, water-column data (e.g., turbidity, 

temperature, salinity, saturation state), and 

benthic substrate type (Davies and Guinotte 

2011). These variables, particularly hard 

substrata, can be patchy at small scales, and 

these alone may determine whether or not 

deep-sea corals inhabit a given location. In 

addition to predictor variables, there are 

limitations with the coral records used in 

models. The taxonomy of deep-sea coral 

records can be highly uncertain. This is 

particularly true for deep-sea coral records 

obtained from video and/or images when 

no specimens were collected for expert 

examination. Coral records obtained from 

trawl/bycatch surveys are associated with 

inherent spatial uncertainty as trawls can 

be several kilometers in length and precise 

georeferencing can be difficult depending on 

the technology used. 

Perhaps the most important question that 

arises pertains to the accuracy of these models 

for deep-sea corals. The preferred method to 

assess model accuracy is to perform robust 

field validation surveys to assess model 

performance. Field validation efforts must 

be conducted to assess the accuracy of the 

models, enable model refinement, and gauge 

the utility of these methods for determining

deep-sea coral habitat in unsurveyed areas,

as illustrated in Figure 1. Rooper et al. (2016)

demonstrated a systematic field validation

for a U.S. deep-sea coral predictive habitat

model for the Eastern Bering Sea, and similar

efforts are currently underway in the Aleutian

Islands, Gulf of Mexico and Northeast

U.S. Field validation of models should be

integrated into the cruise plans of existing/

future NOAA deep-sea exploration and

research. These surveys should record both

georeferenced locations of coral presence

(identified to the best taxonomic resolution

possible) as well as locations of areas where

corals have been confirmed to be absent. They

should also be appropriately stratified so that

areas predicted to have low, intermediate,

and high habitat suitability are sampled

adequately to support model calibration and

validation. These accuracy assessments will

guide refinement of future deep-sea coral

predictive habitat modeling efforts, and also

shape the appropriate use of such models

for conservation and management purposes.

If field validation is not possible, crossvalidation methods can be used to aid in the

assessment of predictive model results (e.g.,

Huff et al. 2013).

VI. Conclusions and

Recommendations

Predictive habitat modeling is increasingly 

used to help identify areas that are most likely 

to harbor deep-sea coral taxa. The growth 

and adoption of these techniques in most U.S. 

regions are primarily due to improvements 

in input data quality/quantity, the low cost 

of producing the models, and the need to 

identify habitat across large spatial extents 

for management. The spatial resolution of 

model results has improved in the last five 

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

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230

years to the point where model outputs 

can be used to aid management decisions, 

and this is also spurring an increase in their 

development and subsequent use. Some of the 

regional examples reviewed here have been 

used to help target areas for field sampling 

efforts and others have been used to help 

inform U.S. regional Fishery Management 

Councils’ EFH/HAPC review processes. Highresolution substrate data (e.g. backscatter, side 

scan sonar) can be used in conjunction with 

model results to identify areas with the highest 

probability of finding deep-sea corals, but the 

limited availability of substrate data at regional 

scales continues to hinder the accuracy of 

predictive habitat models. Model results tend 

to over predict suitable deep-sea coral habitat 

in many regions due to the absence of highquality substrate data and this issue will likely 

persist as the distribution of substrate is highly 

variable over small spatial scales. 

VII. Acknowledgments

Marine Conservation Institute and Bangor 

University would like to thank the Ittleson 

Foundation, Oak Foundation, NOAA’s 

Deep-Sea Coral Research and Technology 

Program, and ESRI’s Conservation Program 

for providing funding and software support 

for modeling efforts. We also would like to 

thank Bob Stone and Chris Rooper for their 

input on Alaskan modeling efforts. BPK 

would like to thank Matt Poti, Chris Caldow, 

Dave Packer, Martha Nizinski, Amy Drohan, 

Richard Langton, Vincent Guida, Peter 

Etnoyer, Andrew David, Thomas Hourigan, 

Gregory Boland, William Shedd, James 

Sinclair, Christopher Jenkins, and Laughlin 

Siceloff for helpful discussions, data and 

technical assistance. BPK was supported 

under NOAA Contract No. DG133C07NC0616 

with Consolidated Safety Services, Inc.

Recommendations to 

NOAA Deep-Sea Coral 

Research and Technology 

Program:

1. Encourage all Program funded cruises

to provide presence and absence data at

relevant spatial scales to help validate and

improve predictive habitat models.

2. Develop and improve the publicly

available national database of coral

occurrences and absences for the U.S. 

EEZ, including in situ environmental data 

where possible.

3. Support the development of a publicly

available database for high-resolution

substrate data for the U.S. EEZ.

4. Ensure that the modeling results funded

by the Program are incorporated into U.S. 

regional Fishery Management Councils’

EFH/HAPC reviews and other relevant

Council processes.

5. Provide support to make the best regional 

predictive habitat model maps broadly

available to the public and federal 

agencies for marine spatial planning

purposes.

6. Convene regular working groups to

compare and synthesize the results of 

modeling efforts in each region. Multiple 

modeling efforts lead to more robust 

understanding, but require expert review,

comparison, and synthesis to identify

strengths and weaknesses of different

approaches as well as areas of consensus

and disagreement among models.

7. Ensure that field surveys, habitat

modeling, and management efforts are

efficiently integrated to take advantage of

the feedback loops illustrated in Figure 1.

8. Improve mapping and modeling of

real and potential human impacts and

integrate these maps with deep-sea coral 

PHM to identify and prioritize high risk 

areas.

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231

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Rengstorf AM, Grehan A, Yesson C, Brown 

C (2012) Towards high-resolution habitat 

suitability modeling of vulnerable marine 

ecosystems in the deep-sea: Resolving terrain 

attribute dependencies. Marine Geodesy 

35:343–361

Rengstorf AM, Yesson C, Brown C, Grehan AJ 

(2013) High-resolution habitat suitability 

modelling can improve conservation of 

vulnerable marine ecosystems in the deep-sea. 

Journal of Biogeography 40:1702–1714

Rooper CN, Sigler M, Goddard P, Malecha 

PW, Towler R, Williams K, Wilborn R, 

Zimmermann M (2016) Validation and 

improvement of species distribution models for 

structure forming invertebrates in the eastern 

Bering Sea with an independent survey. Mar 

Ecol Prog Ser 551:117-130

Hermann AJ (2014) Predictive models of 

coral and sponge distribution, abundance 

and diversity in bottom trawl surveys of the 

Aleutian Islands, Alaska. Mar Ecol Prog Ser 

503:157–176

Rooper CN, Zimmermann M, Prescott MM (2017) 

Comparison of modeling methods to predict 

the spatial distribution of deep-sea coral 

and sponge in the Gulf of Alaska. Deep Sea 

Research I 126:148-161

Ross RE, Howell KL (2013) Use of predictive 

habitat modeling to assess the distribution 

and extent of the current protection of ‘listed’ 

deep-sea habitats. Diversity and Distributions 

19:433–445

Ross SW, Brooke S (2012) Research cruises. 

Deepwater canyons 2012: pathways to 

the abyss. Legs I and II. NOAA Office 

of Exploration and Research, Bureau 

of Ocean Energy Management, and US 

Geological Survey. Additional information 

available at: http://oceanexplorer.noaa.gov/

explorations/12midatlantic/background/

habitats/habitats.html and http://

deepwatercanyons.wordpress.com

SAFMC (2013) Decision Document for Coral 

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for Coral, Coral Reef, and Live/Hardbottom 

Habitats in the South Atlantic Region. Available 

at: http://safmc.net/useful-info/coralamendment-8/

Scanlon KM, Waller RG, Sirotek AR, Knisel JM, 

O’Malley JJ, Alesandrini S (2010) USGS coldwater coral geographic database - Gulf of 

Mexico and Western North Atlantic Ocean, 

Version 1.0. USGS Open File Report 2008-1351. 

US Geological Survey, Reston, VA. Available at 

http://pubs.usgs.gov/of/2008/1351/

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Stone RP, Guinotte JM, Cohen AL, A. H, Cairns SD 

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Stone RP, Shotwell SK (2007) State of Deep Coral 

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Lumsden SE, Hourigan TF, Bruckner AW, Dorr 

G (eds) The State of Deep Coral Ecosystems of 

the United States. Department of Commerce, 

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40:1688–1701

Tittensor DP, Baco AR, Brewin PE, Clark MR, 

Consalvey M and others (2009) Predicting global 

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Tong R, Purser A, Guinan J, Unnithan V (2013) 

Modeling the habitat suitability for deep-water 

gorgonian corals based on terrain variables. 

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Tracey DM, Rowden AA, Mackay KA, Compton T

(2011) Habitat-forming cold-water corals show 

affinity for seamounts in the New Zealand 

region. Mar Ecol Prog Ser 430:1–22

White M, Mohn C, Orren MJ (2007) Physical 

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Pitcher TJ, Morato T, Hart PJB (eds) Seamounts: 

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Wilson MFJ, O’Connell B, Brown C, Guinan JC, 

Grehan AJ (2007) Multiscale terrain analysis 

of multibeam bathymetry data for habitat 

mapping on the Continental Slope. Marine 

Geodesy 30:3–35

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modeling of coral distribution in the Central 

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(2017). The global distribution of deep-water 

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suitability of cold water octocorals. Journal of 

Biogeography 39(7):1278–92

PREDICTIVE HABITAT MODELING FOR DEEP-SEA CORALS IN U.S. WATERS

• • •
             

             

           

           

     

An octopus hides in a rock wall

dotted with cup coral and soft coral

in Welker Canyon off New England.

Courtesy of the NOAA Office of

Ocean Exploration and Research.
                     

     

                     

                 

                 

                     

               

                   

                         

                     

                     

                       

                     

                     

                 

                     

                       

                     

                     

                   

               

                   

                   

                     

                      

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

STATE OF DEEP-SEA 

CORAL AND SPONGE 

ECOSYSTEMS OF THE 

NORTHEAST UNITED 

STATES 

David B. 

Packer1*, 

Martha S. 

Nizinski2, 

Michelle S. 

Bachman3, 

Amy F. 

Drohan1, 

Matthew Poti4, 

and Brian P. 

Kinlan4

  

1 NOAA Habitat Ecology

Branch, Northeast

Fisheries Science Center,

Sandy Hook, NJ  

* Corresponding Author:

Dave.Packer@noaa.gov

  

2 NOAA National

Systematics Laboratory

Smithsonian Institution,

Washington, DC

3 New England Fishery

Management Council,

Newburyport, MA

4 NOAA Marine Spatial

Ecology Division, National

Centers for Coastal Ocean

Science, Silver Spring, MD

I. Introduction

The Northeast region extends from Maine to North Carolina ends at

the U.S. Exclusive Economic Zone (EEZ). It encompasses the

continental shelf and slope of Georges Bank, southern New

England, and the Mid‐Atlantic Bight to Cape Hatteras as well as

four New England Seamounts (Bear, Physalia, Mytilus, and

Retriever) located off the continental shelf near Georges Bank (Fig.

1). Of particular interest in the region is the Gulf of Maine, a semi‐

enclosed, separate “sea within a sea” bounded by the Scotian Shelf

to the north (U.S. jurisdiction ends at the Hague Line), Georges

Bank to the south, and Cape Cod to the southwest. Diverse benthic

habitats are found on the rocky ledges, soft sediment banks, and

within the 21 deep basins (the largest and deepest being Jordan,

Wilkinson, and Georges) of the Gulf. Georges Bank, another

prominent feature in the region, is a shallow elongate extension of

the continental shelf bounded by the Gulf of Maine to the north,

Great South Channel and Nantucket Shoals to the west and south,

and the Northeast Channel/Scotian Shelf to the east and north (U.S.

jurisdiction ending at the Hague Line). The central portion is

shallow, consisting mostly of larger grain‐sized sediments, ranging

from sand to gravel to boulders depending on location. Numerous

submarine canyons, found near the shelf break from Georges Bank

down to Cape Hatteras (Fig. 1), cut into the slope and occasionally

into the shelf. They were formed by erosion of sediments and

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 1. The major submarine canyons and seamounts (within the EEZ) off the Northeast coast of the U.S.

sedimentary rocks of the continental margin

and are classed as deep (V‐shaped from erosion

by rivers, mass wasting, and turbidity currents)

or shallow (shallowly eroded into the

continental margin, or U‐shaped canyons). The

Mid‐Atlantic Bight includes the shelf and slope

from Georges Bank to Cape Hatteras. Many

hard‐bottom topographic features characteristic

of other deep‐sea coral habitats (e.g., boulders,

bedrock, etc.) are absent here. The relatively

small amount of hard substrate in deeper water

available in this area occurs in conjunction with

submarine canyons or is anthropogenic in

origin (e.g., artificial reefs, shipwrecks). The

shelf topography is relatively smooth and

sediments are rather uniform, mostly consisting

of sand with some gravel, with finer sediments

occurring toward the outer shelf, shelf break,

and deeper parts of canyons. The Hudson Shelf

Valley and Canyon, off NY/NJ is the main

physiographic feature in the Bight, extending

from the inner shelf to slope.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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Packer et al. (2007) extensively reviewed most

of the major coral groups of the Northeast (the

exception was sea pens or pennatulaceans; see

below), including gorgonians and true soft

corals (alcyonaceans), black corals

(antipatharians), and certain stony corals

(scleractinians). However, much of the

information needed to assess the status of the

deep‐sea coral habitats was unavailable or

incomplete at the time of that review. There

was missing or inadequate information on

deep‐sea coral distribution, abundance, natural

history, taxonomy, and overall population

status and trends, as well as a lack of high‐

resolution bathymetry data. Available

information on deep‐sea corals from shelf,

slope, and canyon locations was compiled from

historical databases (see Packer et al. 2007).

These data were mainly presence‐only or

occurrence records. The canyons where the

majority of the survey work had been done

were Heezen, Lydonia, Oceanographer,

Hendrickson, Baltimore, and Norfolk canyons.

A few patterns or trends were suggested or

apparent (Watling and Auster 2005; Packer et

al. 2007). For the gorgonians and true soft

corals, species composition varies with depth.

The suite of species that occurs at depths less

than 500 m (shelf and upper slope) is different

than that which occurs at depths greater than

500 m (lower slope and rise) (Watling and

Auster 2005). Some other species (e.g., the stony

coral Astrangia poculata) can also be found in

nearshore waters less than 50 m deep. Corals

occurred in higher densities and were more

diverse in canyons, and some species, such as

those restricted to hard substrates, were found

only in canyons while those utilizing soft

substrata were found both in canyons and on

the continental slope (see discussions of Hecker

and colleagues surveys in Packer et al. 2007).

Surveys conducted prior to 2012 (Moore,

Watling, Auster, Shank, and colleagues)

showed that deep‐sea corals are major

components of benthic communities of the New

England Seamounts. At the time, the black

corals (antipatharians) in this region appeared

to be primarily confined to the seamounts. Sea

pens (pennatulaceans) were not discussed in

detail. However, the most common and

widespread sea pens (Pennatula aculeata and

Stylatula elegans) are found on the continental

shelf in soft sediments, with most other species

recorded on the slope and in canyons.

In addition, little was known about the fauna

associated with deep‐sea coral habitats. For

example, debate continued on the role of deep‐

sea corals as essential fish habitat (EFH) in this

region. Although redfish (Sebastes spp.)

commonly occur in the vicinity of deep‐sea

corals in the Gulf of Maine and corals may

provide important structural attributes of

habitat that can affect the distribution and

abundance of fishes, it was suggested that

corals may not be functionally different from

other structure available to fishes (e.g., Auster

2005). There was also little information

available about associations between

invertebrate species and deep‐sea corals in this

region, except for nearby studies in Canada

(e.g., Northeast Channel: Metaxas and Davis

2005). Studies on invertebrate species

associations with octocorals on the seamounts

were ongoing.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

The effects of fishing efforts on deep‐sea corals

and coral habitats in this region were not

quantified. Quantitative information on the

extent of other anthropogenic impacts to corals

was also not available. The types of bottom‐

contact fishing gear used in the Northeast

include fixed gear such as longlines, gillnets,

pots and traps, as well as mobile gears such as

trawls and dredges. Most fishing is conducted

on the shelf, or along the shelf break, and in the

Gulf of Maine.

The fisheries that have the highest likelihood of

occurring near concentrations of known deep‐

sea coral habitats in canyon and slope areas are

the bottom trawl fishery for monkfish (or

goosefish, Lophius americanus) and the bottom

longline fishery for tilefish (Lopholatilus

chamaeleonticeps, Caulolatilus microps), and the

pot fisheries for Atlantic deep‐sea red crab

(Chaceon quinquedens) and offshore lobster

(Homarus americanus). Bottom trawl fisheries for

squid (Doryteuthis [Amerigo] pealeii and Illex

illecebrosus), whiting (includes silver hake,

Merluccius bilinearis and offshore hake,

Merluccius albidus), and butterfish (Peprilus

triacanthus), and pot fisheries for Jonah crab

(Cancer borealis) occur along the shelf break. In

the Gulf of Maine, lobster fishing occurs in all

known coral habitats, and gillnet and bottom

trawl fisheries for groundfish such as pollock

(Pollachius virens), redfish (Sebastes spp.), and

white hake (Urophysis tenuis) occur offshore.

In terms of fishery management actions, in

2005, NOAA’s National Marine Fisheries

Service (NMFS), on the advice of the New

England and Mid‐Atlantic Fishery Management

Councils (NEFMC, MAFMC), closed portions

of Oceanographer and Lydonia canyons

indefinitely to vessels fishing with bottom

trawls or bottom gillnets during a monkfish

“day‐at‐sea” in order to minimize impacts of

the monkfish fishery on EFH. Although corals

were not included in the EFH descriptions for

any fisheries species in the Northeast region, a

number of federally‐managed species had EFH

defined as hard substrates in depths greater

than 200 m (e.g., redfish), and the abundance of

structure‐forming organisms, especially deep‐

sea corals in these canyons was a major driver

for the protection of these hard bottom habitats.

In this chapter we highlight new scientific

information on deep‐sea coral ecosystems that

has become available since the publication of

Packer et al. (2007), both from the analysis of

samples collected before 2007 (e.g., seamount

expeditions conducted from 2003‐2005 and

described in Packer et al. 2007) and major new

mapping and research initiatives since 2011.

This new information has resulted in major new

management measures established or under

development to help protect these habitats in

the region. We also include information on

deep‐sea sponges, which were not addressed in

the 2007 report.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

II. Update on Scientific

Information

II.1. New Research – Overview

In 2010, the Bureau of Ocean Energy

Management (BOEM), in partnership with the

U.S. Geological Survey (USGS) and the

National Oceanic and Atmospheric

Administration (NOAA) initiated a major

study: Exploration and Research of Mid‐

Atlantic Deepwater Hard Bottom Habitats and

Shipwrecks with Emphasis on Canyons and

Coral Communities (BOEM 2010). Between

2012 and 2015, NOAA conducted a series of

exploratory surveys of coral habitats in the Gulf

of Maine, additional Northeast canyons, and

New England seamounts, which have

significantly expanded our knowledge of deep‐

sea coral and sponge habitats in the region.

While analysis of information from many of

these surveys is ongoing, several publications

and reports have already come out (cited and

discussed below), and the higher‐level

identification of deep‐sea coral and sponge

habitats has already been used by the Regional

Fishery Management Councils in development

of coral protection zones and by the Obama

Administration to inform designation of the

Northeast Canyons and Seamounts Marine

National Monument (discussed below).

II.1.i – BOEM Atlantic Deep-water

Canyons Study 2010-2017

This multi‐year study focused on Baltimore and

Norfolk canyons in the southern mid‐Atlantic

Bight, and was designed to understand the

distribution and complexity of hard bottom

communities in the region. The study was

funded by BOEM and awarded to Continental

Shelf Associates, Inc. Partners included NOAA,

USGS, North Carolina Museum of Natural

Sciences, European agencies, and several U.S.

universities. The study area was originally

selected based on the potential oil and gas lease

sales in federal waters within the BOEM mid‐

Atlantic planning region.

Specific coral related objectives included

characterization of the canyons’ physical

environments, habitats, and associated

communities, and included documenting deep‐

sea coral distribution and diversity, developing

predictive habitat models of deep‐sea coral

distributions, evaluating sensitivity of

communities to disturbance, and

understanding population connectivity

between canyons. The study mapped

Baltimore, Accomac, Washington, and Norfolk

canyons, with subsequent remotely operated

vehicle (ROV) surveys, collections, and studies

focused on Baltimore canyon and Norfolk

canyon.

For example, see Brooke et al. 2017 for deep‐sea

corals specifically, as well as other studies by

Ross et al. 2015; Prouty et al. 2016; Bourque et

al. 2017, etc. See also:

http://oceanexplorer.noaa.gov/explorations/12

midatlantic/welcome.html and

http://oceanexplorer.noaa.gov/explorations/13

midatlantic/welcome.html.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 2. ACUMEN priority areas selected by NOAA and external partners. Mapping efforts by NOAA Ship

Okeanos Explorer depicted as color swaths; NOAA Ship Ferdinand Hassler as yellow boxes.

II.1.ii – ACUMEN (Atlantic Canyons

Undersea Mapping Expeditions) 2012

Because the submarine canyons on the

Northeast continental shelf and slope are

diverse and unique habitats that contain a

variety of fauna, they are of great interest to

federal and state management agencies.

However, most are poorly known, due to the

costs and logistical difficulties of surveying

these areas. A partnership among NOAA line

offices, the Atlantic Canyons Undersea

Mapping Expeditions (ACUMEN) was

established to make efficient use of research

ships’ multibeam mapping resources, allowing

for effective and efficient data collection and

capitalizing on complementary capabilities of

NOAA assets to produce an integrated,

coherent dataset. ACUMEN’s goals included

field efforts to support the NOAA Habitat

Blueprint Northeast Regional Initiative (NOAA

2012a), support of NOAA Integrated Ocean and

Coastal Mapping efforts, prioritization of

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

canyons for conservation goals as suggested by

state and regional constituents (e.g., NEFMC

Habitat Plan Development Team [Habitat PDT],

the Mid‐Atlantic Regional Council on the

Ocean [MARCO]), and sharing of data and

products across platforms to guide and refine

expedition plans in near real‐time. A working

relationship was also established with scientists

in charge of the BOEM Atlantic Deep‐water

Canyons study mentioned above.

Priority areas along the continental shelf/slope

from Virginia to Rhode Island were initially

identified for exploration and mapping by

ACUMEN to gather baseline information in

support of science and management needs.

Between February and August 2012, five

expeditions conducted a mapping ‘blitz’

focused on the submarine canyons (Fig. 2). The

expeditions highlighted the complementary

capabilities of three NOAA ships: Okeanos

Explorer (Office of Ocean Exploration and

Research), Ferdinand R. Hassler (Office of Coast

Survey), and Henry B. Bigelow (NMFS,

Northeast Fisheries Science Center). The Hassler

and Okeanos Explorer collected high‐resolution

bathymetry data that were quickly processed

into mapping products that guided the majority

of subsequent habitat modeling and deep‐water

surveys (discussed below).

The 2012 ACUMEN field efforts finished with a

July survey aboard the FSV Henry B. Bigelow.

Overall goals of the mission were to survey and

ground‐truth known or suspected deep‐sea

coral habitats associated with the submarine

canyons off the edge of the Northeastern U.S

continental shelf/slope. Activities included: (1)

characterizing benthic habitats and identifying

areas where deep‐sea corals and sponges were

present; (2) initial ground‐truthing of areas

predicted to be coral “hotspots” based on data

and outputs from a deep‐sea coral habitat

suitability model; and, (3) ground‐truthing

newly collected high‐resolution (25‐50 m)

continental slope bathymetric maps created

from multibeam data collected during

ACUMEN cruises. Using the Woods Hole

Oceanographic Institution’s (WHOI) towed

camera system (TowCam), three main canyon

areas were targeted: Toms Canyon complex in

the mid‐Atlantic (Toms, Middle Toms, and

Henderickson canyons; Toms‐Hendrickson

inter‐canyon area), Veatch Canyon and the rim

of an unnamed canyon northeast of Veatch, and

Gilbert Canyon off New England (Fig. 3).

Gilbert Canyon in particular was identified as a

deep‐sea coral “hotspot” by the habitat

suitability model. All three main canyon areas

were either under‐explored or unexplored with

regards to deep‐sea coral and sponge

occurrences. Thus, these areas were of

particular interest to the regional Fishery

Management Councils.

II.1.iii – Physalia Seamount Expedition 2012

In October 2012, the Waitt Institute funded a

cruise with their two REMUS 6000 autonomous

underwater vehicles (AUVs; operated by

Woods Hole Oceanographic Institution to

investigate deep‐sea corals on Physalia

Seamount (summit depth approximately 1880

m), a previously unexplored member of the

western New England Seamount chain within

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                            236

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

New

Figure 3. Map of 2012‐15 NOAA expeditions (dives and tows) of the continental slope, canyons, and

Mytilus, Physalia, and Retriever Seamounts using either the Woods Hole Oceanographic Institute’s

TowCam towed camera system or the Canadian ROV ROPOS onboard NOAA ship Henry Bigelow, and

exploratory surveys by ROV Deep Discoverer onboard NOAA ship Okeanos Explorer. BOEM funded

surveys of Baltimore and Norfolk canyons are not included. A) England region. B) Mid‐Atlantic region.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

the U.S. EEZ (Kilgour et al. 2014; Fig. 3). The

vehicles collected 2956 color seafloor images as

well as 120 kHz (low‐frequency) and 420 kHz

(high‐frequency) sidescan sonar. The cruise

demonstrated that AUVs are suitable for the

rapid assessment of the presence and

distribution of deep‐sea corals. This ʺhigh and

fastʺ sampling strategy was appropriate for the

spatial resolution of current management

approaches that essentially propose broad

geomorphic features (e.g., discrete submarine

canyons, seamounts) as deep‐sea coral

protection zones. The geo‐referenced images

provided fine scale distribution information,

which is optimal for surveying the large areas

required for tactical management needs

(Kilgour et al. 2014). This project demonstrated

that AUVs could be ideal tools for linking fine‐

scale spatial distribution of deep‐sea corals to

meso‐scale patterns and variation in landscape

features, flow regimes and other oceanographic

attributes. Such information is of particular

benefit for improved deep‐sea coral habitat

suitability modeling to predict distribution of

deep‐sea coral taxa across regional landscapes

(Kilgour et al. 2014). The results from the AUV

deep‐sea coral surveys at Physalia Seamount

were provided to the NEFMC for use in their

regional deep‐sea coral management zone

determinations.

II.1.iv – NOAA Ship Okeanos Explorer

Atlantic Canyon and Seamounts

Expeditions 2013 & 2014

In 2013 and 2014, NOAA Ship Okeanos Explorer

surveyed canyons off New England and the

Mid‐Atlantic as well as three seamounts off

New England using the ROV Deep Discoverer

(Fig 3). Areas for exploration were prioritized

prior to the cruise by resource managers,

federal and state partners, and through broad

participation of the marine science community.

The ship also conducted additional multibeam

mapping.

In 2013, surveys were conducted in 11

submarine canyons and Mytilus Seamount

(Quattrini et al. 2015; also:

http://oceanexplorer.noaa.gov/okeanos/explorat

ions/ex1304/welcome.html). The New England

areas surveyed in 2014 included Physalia and

Retriever Seamounts, Nantucket Canyon, an

unnamed, minor canyon east of Veatch

Canyon. Mid‐Atlantic areas surveyed in 2014

included the Mey‐Lindenkohl Slope

(Lindenkohl and Hendrickson canyons) and

Washington, Norfolk, Phoenix, McMaster, and

Ryan canyons. Full descriptions of the 2014

dives can be found at:

http://oceanexplorer.noaa.gov/okeanos/explorat

ions/ex1404/welcome.html.

II.1.v – NOAA Northeast Deep-sea Coral

and Sponge Initiative 2013-2015

The Deep Sea Coral Research and Technology

Program funded a Northeast fieldwork

initiative from 2013 to 2015 to locate, survey,

and characterize deep‐sea coral and sponge

communities in this region (NOAA 2016). The

fieldwork initiative was implemented by

NOAA scientists in collaboration with other

government agencies (including the Canadian

Department of Fisheries and Oceans), and

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

researchers from academic institutions. The

projects were designed with the participation of

the Regional Fishery Management Councils.

The emphasis during the 3‐year initiative was

on baseline coral and sponge habitat

characterization, while trying to survey as

many areas as possible in order to provide

information and contribute to management

decisions by the Fishery Management Councils

and other regional partners (e.g., Mid‐Atlantic

Regional Council on the Ocean, [MARCO]).

Thus, the Initiative was designed to satisfy

resource management needs while significantly

increasing our understanding of the region’s

deep‐sea coral and sponge ecosystems.

By combining Program resources with other

partners within and outside of NOAA,

leveraging funding, and employing a wide

range of research tools, the initiative advanced

deep‐sea coral science and management

through three major fieldwork projects:

1. Exploratory surveys of coral/sponge

habitats in submarine canyons, slope areas,

and seamounts off New England and the

Mid‐Atlantic (Fig 3). Surveys off New

England and the Mid‐Atlantic were

conducted every summer from 2013‐2015

using towed cameras, targeting areas in and

around submarine canyons from

approximately 300+ m to 2100 m. The Deep

Sea Coral Research and Technology

Program was also a partner on the Okeanos

Explorer expeditions. Surveys contributed to

groundtruthing and refining the next

iterations of the Northeast’s deep‐sea coral

habitat suitability model.

2. Characterizations of seafloor communities

in the U.S. and Canadian cross‐boundary

Gulf of Maine region and on the U.S. and

Canadian continental margin (ROV ROPOS

dives, 2014; Fig 3).

3. Exploratory surveys of northern Gulf of

Maine (U.S.) habitat areas for deep‐sea

corals and sponges. This component also

included collecting specimens of the

common sea pen (Pennatula aculeata) to

determine if they are being used by fish

larvae (perhaps redfish, Sebastes spp.) as

nursery habitat, as has been observed in

Canada (Baillon et al. 2012).

II.2. Taxonomy and Species

Distributions

II.2.i – Corals

a. Coral taxonomy

Deep‐sea corals in the Northeastern U.S. belong

to four orders: Order Scleractinia (the stony

corals), Order Antipatharia (black corals),

Order Alcyonacea (including gorgonians

[formerly in the order Gorgonacea], true soft

corals, and stoloniferan corals), and Order

Pennatulacea (sea pens) (see Packer et al. 2017

online). Sea pens were not covered in detail by

Packer et al. 2007; however, they may play an

important role in benthic communities (Buhl‐

Mortensen et al. 2016) and provide nursery

habitat for some commercial fish species (see

below, re: Baillon et al. 2012) Thus, sea pens are

now included in the species list and summaries

of deep‐sea corals in this region. NOTE:

Gorgonians are the most important structure‐

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b. Coral distribution

           

               

             

                 

                

                                   

                               

                           

                                 

           

              

     

     

               

           

     

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Table 1. Number of coral species reported from the Northeast U.S. in 2007 and in the present report

(including taxa with incomplete taxonomy – e.g. taxa reported only to genus level)). Increases include new

species identifications and range extensions, but principally represent species not included in the first report.

The lower number of sea pen taxa reflects a number of synonymized species. Data sources and references

online in Packer et al. 2017: https://deepseacoraldata.noaa.gov/library/2017‐state‐of‐deep‐sea‐corals‐report.

.TAXON Packer et al.

2007

Packer et al.

2017

Black corals (Order Antipatharia) 2 6

Stony corals (Order Scleractinia) 16 17

Gorgonian corals (Order Alcyonacea – in part [formerly

Order Gorgonacea])

17 32

True soft corals and stoloniferans (Order Alcyonacea) 9 9

Sea pens (Order Pennatulacea) 17 14

Total 61 78

forming corals in the Northeast region, and for

the purposes of this chapter, they will generally

be treated separately from other alcyonaceans.

Current taxonomic recognizes gorgonians, true

soft corals, and stoloniferans under order

Alcyonacea (Bayer 1981; Daly et al. 2007;

McFadden et al. 2010).

Packer et al. (2007) reported 61 species of corals

from the Northeast region (Table 1). Since 2007,

the systematic compilation of coral records in

the National Deep‐sea Coral and Sponge

Database, along with new descriptions and

range extensions, has increased this number to

78 (Table 1; see Packer et al. 2017 online). The

largest increase in numbers was among the

gorgonian corals, reflecting new species

descriptions by Cairns (2006, 2007), Pante and

Watling (2011), and Simpson and Watling

(2011), as well as new additions to the species

list. Many of the new species descriptions

resulted from analysis of specimens collected

during 2003‐2005 seamount expeditions

(described in Packer et al. 2007).

For sea pens, there have been several new

additions and name changes; e.g., Protoptilum

aberrans Kölliker 1880 has been removed from

the current list as it is now considered a junior

synonym of Protoptilum carpenteri Kölliker, 1872

(Mastrototaro et al. 2015), Funiculina armata

Verrill, 1879 has been added, etc. (see Packer et

al. 2017 online).

Data Sources: The primary sources of historical

(i.e., prior to 2007) deep‐sea coral records and

observations in this region are discussed and

referenced in Packer et al. (2007). There are two

main types of deep‐sea coral data in the

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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Northeast region: geo‐referenced presence

records, and presence records with no

accompanying geo‐referenced information, or

only general location information, which we

call “observations.” (The latter most often

comes from the historical deep‐sea coral survey

literature.) There is too little deep‐sea coral

density or abundance data to be useful for

either scientific or management purposes,

especially in aggregate with other occurrence

data. A database of Northeast deep‐sea corals,

based largely on historical geo‐referenced

presence records from the late 1800s to the

present, was updated by incorporating

taxonomic changes made between 2007 and

2013 and adding “new” presence records

gleaned from museum collection databases

(e.g., the Smithsonian Institution’s National

Museum of Natural History collection, which

includes records of coral taxa collected from

various research surveys/expeditions from 1873

through to the present), other data mining

activities, the literature, and new records from

expeditions in the 2000s (e.g., the NOAA‐Ocean

Explorer 2001 “Deep East” expeditions to

Oceanographer, Hydrographer, and Hudson

Canyons and 2003 ʺMountains in the Seaʺ 

expeditions to the New England Seamounts).

Additional records of sea pens (especially

Pennatula aculeata) collected from 1956‐1984

were compiled from various sources (e.g.,

Langton et al. 1990). Records of new species of

octocorals, mostly from Bear and Retriever

Seamounts with some from the submarine

canyons off New England (e.g., Thouarella

grasshoffi Cairns 2006 from Bear Seamount and

Oceanographer Canyon) were obtained from

recently published literature (Cairns et al. 2007;

Thoma et al. 2009; Pante and Watling 2011;

Watling et al. 2011). New records of

antipatharians were also obtained from recently

published seamount literature (Thoma et al.

2009). These records have now been

incorporated into NOAA’s National Database

on Deep‐Sea Corals and Sponges

(www.deepseacoraldata.noaa.gov; Hourigan et

al. 2015).

Deep‐sea Coral Habitat Suitability Modeling:

Because of the prohibitive costs and logistical

difficulties of surveying for deep‐sea corals and

their associated habitats, geo‐referenced deep‐

sea coral location data are often limited and

patchy, and are mostly of presence‐only. This

makes it difficult for resource managers trying

to identify and implement deep‐sea coral

habitat management areas to develop spatially

contiguous measures for coral conservation in

the absence of comprehensive surveys.

Habitat modeling for deep‐sea corals has

become a cost effective approach to predict

locations of deep‐sea corals and other benthic

species, and aid managers in developing deep‐

sea coral management zone alternatives

(Leverette and Metaxas 2005; Bryan and

Metaxas 2007; Davies et al. 2008; Tittensor et al.

2009; Davies and Guinotte 2011; Guinotte and

Davies 2012; Yesson et al. 2012; Vierod et al.

2013). Habitat suitability modeling examines

the associations between the presence and/or

absence of organisms and their relevant

environmental or habitat variables such as

depth, slope, temperature, and related

measures.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

NOAA developed a set of deep‐sea coral

predictive habitat models for this region for a

set of taxonomic groups (Kinlan et al. 2013;

Kinlan et al., in review). The spatial domain of

the models includes the continental shelf and

canyons in New England and the Mid‐Atlantic.

Seamounts are not included, because the model

is based on the footprint of the coastal relief

digital elevation model, which does not include

the seamounts. A machine‐learning technique

called Maximum Entropy modeling, or MaxEnt,

was used to predict suitability of unexplored

habitats based on locations and environmental

characteristics of known deep‐sea coral

presence (Guinotte et al., this volume).

The MaxEnt models were run with selected

predictor (environmental) variables and

presence data for four taxonomic orders of

deep‐sea corals represented in the Northeast

historical database (Alcyonacea, [formerly]

Gorgonacea, Scleractinia, and Pennatulacea).

Data included in the models were: 1) Northeast

coral presence records from the Cold Water

Coral Geographic (CoWCoG) database (Scanlon

et al. 2010; NOAA 2011a) supplemented with

additional records from the region (Packer and

Dorfman 2012); 2) high‐resolution bathymetry

data from the NOAA Coastal Relief Model

(NOAA 2011b); and, 3) environmental

predictors (seafloor terrain statistics; physical,

chemical, and biological oceanographic data,

and sediment/substrate information).

Habitat suitability maps and model evaluation

methods predicted suitable habitat in the

vicinity of known deep‐sea coral presence

locations, as well as in unsurveyed areas (for

more detailed discussion of model results, see

Guinotte et al. 2016, Guinotte et al., this volume,

Kinlan et al., in review). Some model outputs

are better predictors of coral presence than

others, in part due to the number of coral

records for each order in the historical database.

The model outputs for alcyonacean

(= gorgonian and true soft) corals is based on a

sizeable number of data points and is the model

with the best predictive ability. Therefore, these

model outputs were used by the MAFMC and

NEFMC to evaluate deep‐sea coral habitat

suitability of each proposed broad and discrete

deep‐sea coral management zone (see

discussion, below). Conversely, the model for

scleractinians is based on a smaller number of

records, mostly of solitary, soft‐sediment

dwelling cup corals (e.g., Dasmosmilia lymani

and Flabellum spp.). The scleractinian model is

likely to under‐predict the likelihood of suitable

habitat for this coral taxon. Sea pens were

modeled, based mostly on records from the

continental shelf (e.g., Pennatula aculeata from

New England/Gulf of Maine and Stylatula

elegans from the Mid‐Atlantic) but are not

currently used by the Fishery Management

Councils. These corals occur in soft bottoms, are

more widespread in distribution, and the

contribution of these two species, especially P.

aculeata, as structure‐forming habitat for other

species is still poorly understood or open to

question (see discussion below). Black corals

were not included in the modeling study due to

insufficient data on their distribution when the

study commenced. Future incorporation of

recent data for structure‐forming scleractinians

and black corals in the Northeast region will

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 4a. Example of MaxEnt Predictive Habitat Model (PHM) for gorgonians in the Northeastern U.S.

region (Kinlan et al. 2013). Warm colors (e.g., red) denote areas predicted to be more suitable gorgonian

habitat and cooler coolers (e.g., blue) denote areas predicted to be less suitable gorgonian habitat. Crosses

represent coral presence records used in generating the model.

likely improve the predictive ability of models

for these taxa.  

An example of model outputs for gorgonians

are shown in Fig. 4a,b. The outputs show the

predicted likelihood of deep‐sea coral habitat in

a given area. The predicted likelihood of coral  

habitat suitability is displayed in “thresholded

logistic” maps; i.e., the likelihood values are

displayed by the following likelihood scale:

cooler colors (e.g., blue) denote areas predicted

to be less suitable habitat for gorgonians, while

warmer colors (e.g., red) denote areas predicted  

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 4b. Close‐up of predicted gorgonian habitat suitability in the Oceanographer‐Gilbert‐Lydonia canyon

complex. Crosses represent historical coral presence records. Polygon boundary shows area of interest

identified for exploration and groundtruthing for July 2012 DSCRTP/NEFSC R/V Bigelow cruise.

to be more suitable habitat. The models predict

that the canyons represent “hot‐spots” for

gorgonian coral presence.

After the surveys designed to ground‐truth the

model were conducted (see below), the

Northeast’s habitat suitability models, along

with distribution data from historical and

recent deep‐sea coral surveys, became essential

components of the deep‐sea coral protection

zone decision‐making process by the two

Regional Fishery Management Councils. This

includes the decision by the MAFMC in 2015 to

approve an amendment to protect deep‐sea

corals in their region from the impacts of

bottom‐tending fishing gear along the edge of

the continental shelf and slope and in the

submarine canyons (discussed, below).

Mid‐Atlantic and New England Continental

Slope and Canyons Exploratory Surveys: The

continental slope off the Northeast U.S. is

incised by numerous submarine canyons,

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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which contain extensive hard‐bottom habitat,

which is needed for the attachment of most

deep‐sea corals (Packer et al. 2007). Prior to

2010, most information on deep‐sea coral

habitats within Northeast canyons came from

the surveys of Hecker et al. during the 1980s

(see Packer et al. 2007 for discussions and

references, and B. Hecker presentation in

NEFMC 2011). Based on these historical

surveys, some general patterns in deep‐sea

coral distribution were suggested. As stated

above, deep‐sea corals were generally more

densely distributed and diverse in the canyons

than on the adjacent slope owing to steep

canyon topography, accelerated currents, and

heterogeneous sedimentary geology. Some

species, such as those restricted to hard

substrates, were only found in the canyons

while other species that frequently occur on soft

substrates, such as Acanella arbuscula, were

found both in canyons and on the slope. There

appeared to be two distinct distributional

patterns for the gorgonians and true soft corals

(Watling and Auster 2005). Most were deep‐

water species that occurred at depths > 500 m

(gorgonians in the genera Acanthogorgia,

Acanella, Anthothela, Lepidisis, Radicipes, Swiftia;

the alcyonacean soft coral Anthomastus, and the

stoloniferan Clavularia). Other species appeared

to occur throughout shelf waters to the upper

continental slope (gorgonians Paragorgia arborea,

Primnoa resedaeformis, Paramuricea spp.)

(Watling and Auster 2005). The larger northern

canyons such as Lydonia and Oceanographer

have hard substrates along most of their axes

and walls that support many deep‐sea corals

and sponges. The slope south of Georges Bank

appeared to be mostly soft substrate,

supporting mainly solitary stony (cup) corals

on the upper slope (e.g. Flabellum sp.,

Dasmosmilia lymani) and sea pens deeper than

about 1500 m, with some exceptions (NEFMC

2011). Some harder substrate was found at

depths greater than 1400 m depth on the Mid‐

Atlantic slope off New Jersey. Sparse coral

populations were found at these deeper depths;

sea pens and stony corals appeared to be sparse

or common on the upper slope; sea pens and A.

arbuscula were found on the lower slope. The

larger, southern canyons like Baltimore and

Norfolk canyons appeared to have less hard

substrate than the northern canyons (NEFMC

2011), although some hard substrate that

supports coral and sponges was found in

shallower depths (300‐550 m). There were also

sea pens at upper slope depths and sea pens

and A. arbuscula at lower slope depths.

Recent towed camera and ROV surveys (most

without associated specimen collections) were

conducted in almost all of the major canyons in

the Mid‐Atlantic and New England regions

(Fig. 3). Several minor canyons and slope areas

were also surveyed. The surveys revealed

significant inter‐canyon variability, even

between neighboring canyons. Canyon

morphology, habitat heterogeneity, the amount

and type of substrate, depth, environmental

conditions, and currents appear to play a role in

the biodiversity of corals, sponges, and fishes

observed in each canyon. Corals were observed

in every canyon, however, abundance and

diversity of corals varied dramatically. For

example, only a few colonies of widely

dispersed corals were observed in Washington

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Canyon (off Virginia) whereas a “coral forest”

of the bubblegum coral (Paragorgia arborea) was

observed in Heezen Canyon (near Hague Line).

The surveys also revealed unique areas; e.g.,

discovery of coral biodiversity hotspots in

relatively shallow (300‐500 m) depths at the

head of Wilmington Canyon, as well as

Baltimore and Norfolk canyons (Brooke et al.

2017), and observations of corals in minor

canyons.

Colonial stony corals, the solitary stony cup

coral Desmophyllum dianthus, black corals, most

alcyonaceans, and the majority of sponges

observed were on hard substrates, whereas

other solitary stony corals (e.g., Flabellum sp.),

sea pens, and the bamboo coral Acanella sp.

were common on soft sediments (Fig. 5).

Throughout the Mid‐Atlantic and New

England canyons, gorgonians were the

dominant structure‐forming corals. The large

gorgonians, Paragorgia arborea, Paramuricea spp.

(including P. placomus), Primnoa resedaeformis,

and Acanthogorgia spp. were generally most

abundant in canyons from Virginia to the

Hague Line at depths shallower than 1000 m.

Gorgonians in the genus Anthothela were also

abundant. The bubblegum coral (P. arborea)

reached the largest sizes. As for the true soft

corals, Anthomastus is probably the numerically

dominant species, while Duva florida can be

extremely locally abundant (hundreds of

colonies) but distribution is patchy (Sandra

Brooke, Florida State Univ., pers. comm.). The

first observations of black corals in a number of

the canyons were made during recent surveys.

These included species in the genera

Bathypathes, Parantipathes, Stauropathes, and the

recently described Telopathes magna (Quattrini

et al. 2015; Brooke et al. 2017) (Fig. 5).

Stony corals (Order Scleractinia) were not

especially abundant or diverse at large spatial

scales (although they were patchy and dense

within particular areas of canyons). The solitary

cup coral, Desmophyllum dianthus, was the most

often observed stony coral, usually seen on

canyon walls and in dense aggregations along

ledges or overhangs (Quattrini et al. 2015,

Brooke et al. 2017). The branching colonial

corals Lophelia pertusa and Solenosmilia variabilis

were also observed in the canyons. Both corals

occurred as small, individual colonies, with L.

pertusa generally observed between 300‐1000 m

and S. variabilis generally deeper than 1000 m.

Lophelia pertusa is the most important reef‐

forming deep‐sea coral in the Atlantic, and

forms large reefs (bioherms) in the southeast

U.S. (Ross and Nizinski 2007; Hourigan et al.,

this volume) and the Northeast Atlantic

(Roberts et al. 2009). There was no evidence of

large bioherm formation by either species in the

Northeast region.

Quattrini et al. (2015) reported at least 58 coral

taxa representing 20 families in 11 New

England canyons and adjacent slope habitats.

Quattrini et al. (2015) found that the type of

broad‐scale habitat feature and high habitat

heterogeneity in this region was an important

factor that influenced the diversity of coral

assemblages. There were no significant

differences between deep‐sea coral assemblages

occurring in the two different types of canyons

(continental shelf‐breaching canyons vs.

canyons confined to the continental slope).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 5. Examples of Northeast deep‐sea corals from the submarine canyons and seamounts during the 2012‐

2015 expeditions. Coral identifications in the images may be tentative. a) Various color morphs of gorgonian

bubble gum coral (Paragorgia arborea), Heezen Canyon (656 m) (ROPOS). (b) Black coral Telopathes

magna, Block Canyon (1345 m) (Deep Discoverer). (c) Gorgonian Thouarella grasshoffi (white), soft coral

Anthomastus (red), gorgonian Paramuricea (yellow); Powell Canyon (1300‐1770 m) (TowCam). (d)

Scleractinian colonial coral Lophelia pertusa, Baltimore Canyon (381 m) (Kraken 2). (e) Soft coral

Anthomastus, Oceanographer Canyon (Deep Discoverer). (f). Gorgonian bamboo coral Jasonisis, with

numerous crinoid associates, Mytilus Seamount (Deep Discoverer).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 5 (cont.). Examples of Northeast deep‐sea corals from the submarine canyons and seamounts during

the 2012‐2015 expeditions. Coral identifications in the images may be tentative. g) Two colonies of black corals

and spiraled gorgonian Iridogorgia sp., Retriever Seamount (Deep Discoverer). (h) Stoloniferan coral

Clavularia sp., Nygren Canyon (Deep Discoverer). (i) Scleractinian cup corals and black coral (foreground),

wall of unnamed canyon east of Veatch Canyon (1480 m) (Deep Discoverer). (j) Scleractinian colonial coral

Solenosmilia variabilis and solitary cup coral (above) Desmophyllum dianthus, Baltimore or Norfolk

Canyon (Brooke et al. 2017). (k) Black coral Parantipathes sp., Nygren Canyon (ROPOS). (l). Gorgonian

Paramuricea sp., Welker Canyon (Deep Discoverer).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

However, different, less diverse faunal

assemblages were observed at cold seeps (areas

on the seafloor where hydrocarbon‐rich fluid

seeps up from below) and soft bottom open

slope sites. The canyons often had large patches

of hard bottom deep‐sea coral habitat, which

also included bivalves, anemones, and sponges.

Stony corals (e.g., the solitary cup coral

Desmophyllum dianthus and the colonial

Solenosmilia variabilis) and octocorals were often

abundant on long stretches of canyon walls and

under and around overhangs; the colonial

stony coral Lophelia pertusa was particularly

noted in canyons at depths from 733‐1030 m.

Coral communities were uncommon on the

open slope, except on the channel floor of

Veatch Canyon where sea pens and bamboo

corals in soft sediments were frequently

observed. Corals and sponges were also

observed on boulders and outcrops in some

open slope and inter‐canyon areas. At Veatch

seeps and the canyon wall adjacent to the seep

community in Nygren Canyon, octocorals and

stony cup corals (D. dianthus) were found

attached to authigenic carbonates (carbonate

precipitated in‐situ at the sediment–water

interface and/or within the sediment).

Quattrini et al. (2015) also found that depth was

a significant factor influencing the coral

assemblages. Although species richness did not

change significantly with depth over the range

explored by the surveys (494‐3271 m), species

composition changed at ~1600‐1700 m. Species

composition in the canyons and other areas

with hard substrates were significantly

dissimilar across this depth boundary. Stony

corals and the gorgonians Anthothela spp.,

Keratoisis sp. 1, and Paragorgia arborea, occurred

at depths < 1700 m, whereas gorgonians from

the family Chrysogorgiidae (chrysogorgiids)

and sea pens were more common at depths >

1700 m. Overall, depth, habitat, salinity and

dissolved oxygen explained 71% of the total

variation observed in coral assemblage

structure (Quattrini et al. 2015).

The surveys of both Baltimore and Norfolk

canyons discovered that deep‐sea corals were

patchily distributed and locally abundant and

large gorgonians were the dominant structure‐

forming species on exposed hard substrates

(Brooke et al. 2017). Several species of stony

corals were also present, including new records

of Solenosmilia variabilis on deep walls (~1400 m)

in Norfolk Canyon, and the structure‐forming

Lophelia pertusa on shallower (381‐434 m) walls

of both Baltimore and Norfolk canyons (Brooke

and Ross 2014; Fig. 5d). A new record of the

black coral Telopathes magna was also reported

for Norfolk Canyon. The diversity of structure‐

forming corals was low, but several areas in

both canyons had high coral abundances and

diverse coral communities, dominated by large

octocoral species (Brooke et al. 2017). Coral

distribution varied within and between the two

canyons, with greater abundances of the

bubblegum coral Paragorgia arborea in Baltimore

Canyon than in Norfolk Canyon, but higher

occurrences of stony corals in Norfolk Canyon

than in Baltimore. The gorgonians P. arborea,

Primnoa resedaeformis, and Anthothela grandiflora

had a wide distribution within these canyons,

whereas Paramuricea placomus and the

branching stony corals L. pertusa and S.

variabilis were limited to certain habitat types

and/or depth zones (Brooke et al. 2017).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

The Northeast deep‐sea coral habitat suitability

models generally predict that the canyons are

highly suitable habitat for the gorgonian and

soft coral groups. Surveys of the canyons

allowed independent ground‐truthing of the

models, which were qualitatively validated: all

sites observed to be areas of high coral

abundance and diversity were also areas

predicted to have a high likelihood of suitable

habitat based on the regional model. The

habitat suitability models performed well in

predicting areas of likely deep‐sea coral habitat,

as well as predicting areas where corals are

unlikely to be found. However, the exact

location of areas of high deep‐sea coral density

and diversity often depends on fine‐scale

seabed features (e.g., ridges or ledges of

exposed hard substrate) that are smoothed over

in this regional‐scale model. The current

resolution of the model is grid cells of

approximately 370 m2 (although there are plans

to improve the model by increasing resolution

to 25 m2, as well as incorporating more recent

multibeam bathymetry and coral observations

with higher taxonomic specificity). Habitat

suitability maps based on this model should be

viewed as representing only the general

locations of predicted suitable coral habitat

(within a distance of approximately 350‐740 m,

the dimension of one to two model grid cells).

For this reason, the total area of high/very high

habitat suitability is an approximation using the

best available data. Quantitative ground‐

truthing of the model includes assessment of

the prediction skill of the model and correlation

between predicted habitat suitability and coral

frequency/abundance observations.

New England Seamounts Exploratory Surveys:

Bear Seamount was discussed in Packer et al.

(2007).

Physalia Seamount (summit depth ~ 1,880 m)

was explored by AUVs in 2012 (Kilgour et al.

2014) and an ROV in 2014 (Okeanos Explorer

2014 expedition). The AUV surveys revealed

the presence of octocorals, with sea pens

generally found in flat, soft sediments, and

most other octocorals found at either the

transition between soft sediment and hard

bottom, or on hard bottom features such as

walls, ledges, and gravel/bedrock pavement

(Kilgour et al. 2014). The more detailed ROV

survey revealed relatively low abundances and

low diversity of corals. The gorgonian

Chrysogorgia sp. and sea pen Anthoptilum sp.

were seen most commonly. Of particular note,

Anthoptilum sp. was seen in typical sea pen

habitats embedded in soft sediments but also

on hard substrates; Williams and Alderslade

(2011) described species in this genus from the

Pacific with the unusual adaptation of the

peduncle that acts as a holdfast for attachment

to rocky substrata. Occasionally, the bamboo

coral Lepidisis sp. was observed. Other corals

reported from Physalia Seamount include the

black corals Telopathes sp. and Bathypathes sp.,

the soft coral Anthomastus sp., and stony cup

corals.

Mytilus Seamount has the deepest summit

(~2,500 m) of all New England seamounts in

U.S. waters. The first ROV surveys of Mytilus

(Quattrini et al. 2015) revealed a diverse

assemblage of taxa, including gorgonians (Fig.

5f) and especially black corals, and numerous

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

hexactinellid sponges and demosponges. The

corals observed (> 2600 m) were significantly

different from those observed in Northeast

canyon sites at similar depths by Quattrini et al.

(2015). Differences in species composition

between Mytilus Seamount and other sites were

primarily due to the presence/absence of

numerous species. For example, Chrysogorgia

spp., Convexella? jungerseni, Corallium?

bathyrubrum, Paranarella? watlingi, and

Paragorgia/Sibogagorgia sp. 1 were observed on

Mytilus Seamount, while Acanthogorgia spp.,

Anthothela spp., Clavularia? rudis, P. arborea, and

Paramuricea spp. were not seen on Mytilus

Seamount, but occurred at other sites. No stony

corals were observed here. Quattrini et al.

(2015) suggest that the deeper depths (2600 to

3200 m) are beyond the bathymetric limits of

stony corals.

Retriever Seamount is the farthest‐offshore

seamount within the U.S. EEZ. It is about 2000

m high, 7 km in diameter, and has three main

summits. Thoma et al. (2009) reported the

occurrence of gorgonians in the genera

Chrysogorgia, Iridogorgia, Metallogorgia, Acanella,

and Paramuricea; as well as black corals in the

genera Bathypathes and Parantipathes. A single

ROV dive in 2014 (Fig. 5g) between 2142 to 2003

m depth revealed sandy slopes, boulders and a

rock outcrop. Many sea pen colonies, with

?Anthoptilum sp. more common than Pennatula

sp., as well as stony cup corals (Caryophyllia sp.)

were seen in sedimented areas. Colonies of the

chrysogorgiid gorgonian, Metallogorgia

melanotrichos were very abundant on the rock

outcrops; several “sub‐adult” colonies were

observed, suggesting multiple recruitment

events in the area. Other corals observed on the

outcrop included the gorgonians Hemicorallium

?bathyrubrum (=Corallium ?bathyrubrum) and H.

?niobe, Paramuricea sp., Iridogorgia splendens (at

least one with shrimp associate) and I.

magnispiralis, Candidella imbricata and an

unidentified primnoid coral, bamboo corals

(Lepidisis sp. and Acanella sp.), and black corals

in the genera Parantipathes (branched),

Stauropathes, and, seen further upslope on

isolated rocks, Bathypathes.

Gulf of Maine Exploratory Surveys: Deep‐sea

corals in the Gulf of Maine have been reported

since the 19th century, both as fisheries bycatch

and from naturalist surveys. At one time, deep‐

sea corals may have been considered common

on hard bottoms in the region. However, after a

century of intensive fishing pressure using

mobile bottom gear such as trawls and dredges

as well as fixed gear such as lobster traps, the

denser populations of deep‐sea corals and coral

habitats are now confined to small areas where

the bottom topography makes them mostly

inaccessible to these fisheries (Auster 2005;

Watling and Auster 2005; Cogswell et al. 2009;

Auster et al. 2013).

Previous studies, including work on the

Canadian side of the Gulf (e.g., Northeast

Channel: Buhl and Buhl‐Mortensen 2004;

Watanabe et al. 2009) do show that deep‐sea

corals have a patchy distribution that is

correlated with environmental factors such as

slope, sediment, current, temperature and

depth. Nevertheless, the information needed to

assess their overall status in the U.S. Gulf of

Maine has been lacking. Additionally,

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

information on deep‐sea coral distribution in

relation to habitat and landscape features,

abundance, natural history, associated species,

and human impacts has been inadequate.

Previous deep‐sea coral exploratory surveys

and seafloor mapping in the region guided the

selection of survey sites in 2013. Initial deep‐sea

coral surveys using ROVs in 2002 and 2003

documented a limited number of locations in

Western Jordan Basin and around Mount

Desert Rock with dense coral garden

communities at around 200 m (Auster 2005;

Watling and Auster 2005). Deep‐sea corals were

found on rocks, boulders, ridges and walls

extending above the surrounding fine‐grained

sediments. During a cruise aboard NOAA Ship

Ronald H. Brown in 2005, multibeam bottom

sonar data collected in Western Jordan Basin

revealed that hard bottom in the immediate

area around “114 Bump” (one of the sites

surveyed for corals in 2002‐2003) was more

spatially extensive than previously suspected

(see Fig. 2 in Auster et al. 2014). Thus, the

potential for suitable deep‐sea coral habitat in

the area was more likely.

Results of the recent Gulf of Maine exploratory

surveys (2013‐15) revealed extensive coral

aggregations at depths around 200‐250 m in the

five primary survey sites: Western Jordan

Basin, Central Jordan Basin, near Mount Desert

Rock, on Outer Schoodic Ridge, and on

Lindenkohl Knoll in Georges Basin (Fig. 6)

(Auster et al. 2013, 2014; Packer et al.,

unpublished data). Structure‐forming corals on

hard substrate at all sites were predominantly

gorgonians, although a few tiny, stony

scleractinian cup corals were seen on some

dives. Coral occurrences were classified as

either coral present (sparse to medium density)

or coral garden (high density patches).

Coral gardens are defined as areas where non‐

reef‐forming corals are among the dominant

fauna and occur at densities higher than

surrounding patches (Bullimore et al. 2013).

Dense and extensive gorgonian coral gardens

were seen in Western Jordan Basin, Outer

Schoodic Ridge, and near Mount Desert Rock,

especially in areas of high vertical relief. Outer

Schoodic Ridge especially was a unique area,

with topography reminiscent of narrow slot

canyons on land (e.g., western U.S., in southern

Utah). Based on preliminary analyses of 2013

images (Auster et al. 2013), these steeper areas

had some of the highest densities, with about

16‐39 colonies/m2, well above the threshold of

0.1 colonies/m2 used by ICES (2007) to define

coral garden habitat. Central Jordan Basin and

Georges Basin also contained coral

communities, but these assemblages were more

patchy, less dense, and occurred in lower relief

environments than the aforementioned areas.

The dense corals on the steep vertical walls and

cliffs of Outer Schoodic Ridge and Mount

Desert Rock were primarily Primnoa

resedaeformis, with lower abundances of

Paramuricea placomus (both in two color morphs

of yellow and purple color morphs). On some

of the tall, narrow canyon‐like walls and cliffs

of Outer Schoodic Ridge, P. resedaeformis

colonies were so densely packed it was

impossible to identify and count individual

colonies (Fig. 7a,b), some were likely 1 m tall.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 6. Gulf of Maine deep‐sea coral exploratory survey locations and results (2013‐2015). Modified from

Auster et al. (2014).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Conversely, the major coral species found in

Western and Central Jordan Basin and Georges

Basin was P. placomus, with lower abundances

of P. resedaeformis and Acanthogorgia cf. armata

(Fig. 7c‐e). Paramuricea placomus was found in

higher densities on steeper hard‐bottom areas.

Open areas adjacent to steeper features

including muddy areas containing gravel, sand‐

gravel, and emergent rock outcrop features

(with shallow expressions above the fine‐grain

sediment horizon) supported lower densities of

coral, primarily P. placomus. This was also true

for Outer Schoodic Ridge and Mount Desert

Rock. Based on multivariate analyses of eight

2013 transects in Jordan Basin with coral garden

habitat (Martin 2015), temperature and depth

were primary environmental factors and

sediment type, rock outcrop, and topographic

rise were primary sedimentary factors that

correlated with coral distributions.

Of note were the first observations of Anthothela

(grandiflora?) in the relatively shallow waters of

the Gulf of Maine (Auster et al. 2014). A couple

of colonies were seen at Outer Schoodic Ridge

(Fig. 7f) around 200 m. This species is most

common at 400‐1600 m depth in Northeast

canyons (Hecker et al. 1983; Quattrini 2016;

Brooke et al. 2017) and off the Northeast

Channel along the continental margin

(Cogswell et al. 2009). Paragorgia arborea, both

pink and white forms, which was noted at 114

Bump in Western Jordan Basin during the 2003

survey, was not seen in the more recent

exploratory dives.

The most common and ubiquitous sea pen

species found in the Gulf of Maine, Pennatula

aculeata was also found in dense patches in mud

and gravel/mud habitats adjacent to hard‐

bottom habitats (Fig. 8e,f). Highest densities

were observed in the Mount Desert Rock region

(Packer et al., unpublished data). Pennatula

aculeata became a focus of the surveys because

Baillon et al. (2012) found convincing evidence

that several species of sea pens were directly

utilized as shelter by fish larvae, particularly by

those of redfish (Sebastes spp.; see below, and

Dean et al. 2016).

The results from these Gulf of Maine

exploratory surveys have provided the NEFMC

with improved information to inform

conservation and management decisions for

these unique deep‐sea coral and sponge

habitats. Currently the Council is using these

data to revise and develop new deep‐sea coral

management zone alternatives for the northern

Gulf of Maine region. The large‐sized,

structure‐forming gorgonians P. resedaeformis

and P. placomus, occurring in such high

densities in the relatively shallow waters of the

Gulf of Maine, as compared to the submarine

canyons and seamounts, is unique in the

Northeast. The proximity of these habitats to

shore and their association with commercially

important fish and shellfish increases the

potential role of these habitats as EFH (e.g.,

Auster 2005, 2007). Finding these unique deep‐

sea coral habitats, especially the spectacular

walls of corals at some sites, for the first time

after 40‐plus years of previous underwater

surveys, illustrates how much remains to be

discovered about the Gulf of Maine ecosystem

in order to better conserve and protect its living

marine resources.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 7. Examples of Gulf of Maine deep‐sea coral habitat seen during 2013‐14 exploratory surveys (Auster

et al. 2014 and unpublished images). (a) Dense colonies of Primnoa resedaeformis on a near vertical rock

wall along Outer Schoodic Ridge. (b) Acadian redfish and P. resedaeformis on a near vertical rock wall along

Outer Schoodic Ridge. (c) Purple and yellow forms of Paramuricea placomus and Polymastia sponge along

a steep escarpment in Western Jordan Basin. (d) P. placomus, P. resedaeformis, and juvenile cod in

Western Jordan Basin. (e) P. placomus and haddock, Central Jordan Basin. (f) Anthothela sp. at around 200

m on a near vertical rock wall along Outer Schoodic Ridge.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 8. Examples of dense sponge, anemone, and sea pen habitats in the Gulf of Maine (Auster et al. 2014

and unpublished images). (a) Sponge (including vase sponge Phakellia? sp. and Polymastia sponge sp.) and

anemone habitat along Outer Schoodic Ridge. (b) Sponge (including vase sponge Phakellia?, Polymastia

sponges, and finger sponges) habitat along Outer Schoodic Ridge. (c) Anemone and coral habitat in Western

Jordan Basin. (d) Anemone habitat (with king crab, Lithodes maja) on near vertical rock walls along Outer

Schoodic Ridge. (e) Sea pens, Pennatula aculeata, in soft substrate in Western Jordan Basin. (f) P. aculeata

and Acadian redfish in an area of Jordan Basin known as 91 Bump.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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II.2.ii – Sponges 

Information on the regional sponges and their

habitats is severely lacking. Deep‐sea sponges

in the Northwest Atlantic in general have been

the subject of relatively few studies or surveys.

However, most of these studies, both

historically (e.g., Lambe 1896, 1900) and more

recently (e.g., Fuller 2011; Beazley et al. 2013,

2015, 2016), have been conducted in Canada. In

the Northeast U.S., Theroux and Wigley (1998)

reported on extensive surveys of invertebrates

collected by grab samplers. They reported that

sponges constituted a small proportion of the

observed fauna, and were “generally

uncommon on the continental shelf and

uncommon to rare on the continental slope and

rise.” Larger specimens included Polymastia sp.

and Myxilla sp. usually found on cobbles and

boulders. Sponges were more abundant off

Nova Scotia and in the Gulf of Maine than

elsewhere in the region. However, it is likely

that sponges were underrepresented in their

samples, as benthic grabs cannot adequately

sample hard substrates where most sponges

occur, especially if they are mostly found on

vertical surfaces (e.g., Miller and Etter 2011).

Documenting the diversity and distributions of

sponges in this region is still in the preliminary

stages. Difficulties with taxonomy and

identifications (especially via photographs or

video alone) and lack of expertise hamper these

efforts. Thus far, presence records from the

Smithsonian Institution’s database and other

limited data have been used to create an initial

species list. Most geo‐referenced records that

have been entered in the National Deep‐Sea

Coral and Sponge Database are listed simply as

“Porifera.” New geo‐referenced records of

sponges from the photographic images

obtained from recent 2012‐2015 surveys will be

added to the database after processing.

Sponges that are of conservation interest are the

large, structure‐forming sponges that, like

deep‐sea corals, provide habitat for managed

species of fish (especially juveniles that seek

refuge from predators), increase biodiversity of

local fauna, or are susceptible to fishing gear

impacts (e.g., Auster and Langton 1999; Fuller

2011; Beazley et al. 2013; Maldonado et al.

2016). Preliminary, historical information on the

geographic range (or locations where present),

size, morphological form, and habitats (depth

and substrates) was compiled for 12 structure‐

forming species that are found on the

continental shelf in the region (Table 2).

Encrusting species or species that would not be

considered structure‐forming are not included.

Information sources for Table 2 include the

Marine Life Information Network, the

Stellwagen Bank National Marine Sanctuary

(Gulf of Maine) (on‐line), the European Marine

Life Network, the Marine Life Encyclopedia

website, Georgia Southern University (on‐line),

the Chesapeake Bay Program website, Hartman

(1958), Gosner (1978), Witman and Sebens

(1988), Fuller et al. (1998), Steimle and Zetlin

(2000), and Stokesbury and Harris (2006).

On Georges Bank, species of interest include

Suberites ficus, Haliclona oculata, Halichondria

panicea, Isodictya palmata, Clathria prolifera

(=Microciona prolifera), and Polymastia robusta

(Almeida et al. 2000; Stokesbury and Harris

2006). The larger species are probably the most

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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susceptible to the adverse effects of fishing.

These would include S. ficus, H. oculata, H.

panicea, and I. palmata as well as Mycale lingua

and the large form of Cliona celata. All of these

species attach to some form of hard substrate

including shells. Suberites ficus attaches to small

shell fragments on sandy bottom habitats on

Georges Bank. These relatively rare

microhabitats provide cover for fishes and

crustaceans (Lindholm et al. 2004). As this

sponge grows, the substrate on which it

originally attached is covered over and the

sponge often is rolled along the bottom by

currents and wave action. The other species of

structure‐forming sponges are more common

on hard bottom habitats. Based on the available

information, only two of the species listed in

Table 2, C. celata and H. oculata, are known to

occur south of southern New England (see also

Van Dolah et al. 1987). This may reflect the fact

that natural rocky bottom habitats on the shelf

are rare south of New York Harbor (Steimle

and Zetlin 2000). Other structure‐forming

species of sponge are undoubtedly present in

the Mid‐Atlantic region, but are either found on

hard substrates on the continental slope and in

the canyons or on the shelf attached to gravel,

scallop shells, and shell fragments in

predominantly sandy habitats.

Fuller (2011) also created a species list of

sponges, as documented in published reports,

for the entire Gulf of Maine, including the Bay

of Fundy (Fuller 2011 Appendix III). Sixty‐

seven species were recorded; the Gulf of Maine

Census of Marine Life listing 32 in the Gulf of

Maine. The North Atlantic Fisheries

Organization (NAFO) also published a sponge

identification guide for eastern Canada,

with some overlap into the U.S. Gulf of

Maine (Best et al. 2010).

The species list presented here focuses

mostly on the shallower species. Thus, the

list is preliminary and incomplete. Analysis

of recent deep‐sea surveys will add many

additional species including some new to

science. For example, not listed here are the

vase/basket demosponges (Phakellia/Axinella

spp.). Phakellia ventilabrum has been found

in Stellwagen Bank National Marine

Sanctuary in the Gulf of Maine (< 100 m;

Tamsett et al. 2010), and several large

vase/basket demosponges have been

recently seen in the Gulf of Maine surveys

at around 200 m (Packer et al.

unpublished). However, Fuller (2011) does

not list Phakellia/Axinella as even occurring

in the Gulf of Maine. This type of

conflicting information highlights just how

little is truly known about the deep‐sea

sponges in this region.

Hecker et al. (1980, 1983) indicated that

sponges, along with corals, were frequently the

dominant epifaunal organisms observed during

submersible surveys on exposed hard substrate

in Northeast submarine canyons. Both sponges

and corals were consistently more abundant in

the canyons compared to adjacent slope areas.

The demosponge, Asbestopluma sp. occurred on

walls of consolidated clay in Lydonia Canyon

and accounted for more than half the

invertebrates observed between 800‐950 m

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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Table 2. Preliminary list of structure‐forming sponges of the U.S. Northeast continental shelf.

Species Range Height Form Habitats

Cliona celata Grant,

1826

Gulf of Mexico to

Long Island Sound,

locally to Gulf of St.

Lawrence

Up to 1 m, 60 cm

diameter

Two growth forms, boring

into shells and large

“barrel” shape, firm with

tough outer layer, embeds

rocks and sediments into

tissue

On rock to 200 m; begins life by

boring into limestone, shells, or

calcareous red algae

Halichondria

(Halichondria)

panicea (Pallas,

1766)

Arctic south to Cape

Cod, rarely beyond Up to 30 cm Encrusting, globular, or

branched

Cobbles, boulders, bedrock,

shells, algae down to 80 m (570

m in Europe), esp. abundant in

strong tidal flows

Haliclona

(Haliclona) oculata

(Linnaeus, 1759)

Labrador to Long

Island, rarely to

North Carolina, but

present in Georgia

Up to 45 cm

Short stalk with flat to

rounded finger‐like

branches, very flexible, not

fragile

Sandy, rocky substrates, often

attached to stones, to 150 m

Haliclona

(Haliclona) urceolus

(Rathke and Vahl,

1806)

Range unknown,

found in Bay of

Fundy

To 15 cm, stalk

typically < half body

length

Tubular, even bell shaped,

with thin, hard, flexible

stalk

On rock, shell fragments, etc.

Isodictya

deichmannae (de

Laubenfels, 1949)

Newfoundland to

Rhode Island

Often confused with

I. palmata To 110 m

Isodictya palmata

(Ellis and Solander,

1786)

Nova Scotia to Gulf

of Maine, Georges

Bank

Up to 35 cm Large, palmate with finger‐

like branches

Deep water on rocks, 52‐70 m in

sand and gravel on Georges

Bank

Clathria (Clathria)

prolifera (Ellis and

Solander, 1786)

(=Microciona

prolifera)

Nova Scotia to

Florida and Texas Up to 20 cm

At first encrusting, then

forms small clumps with

fingerlike branches

Shells, pilings, hard surfaces, in

shallow to moderate depths

(52‐70 m on Georges Bank)

Mycale (Mycale)

lingua (Bowerbank,

1866)

Range unknown,

found in the Gulf of

Maine

Up to 30 cm high

with variable width

and depth

In mounds, sometimes in

erect, flattened form with

base narrower than apex

Between 30‐2460 m on rocky

bottom

Myxilla (Myxilla)

fimbriata

(Bowerbank, 1866)

Range unknown,

found in GOM Mounds To 78 m

Polymastia

boletiformis

(Lamarck, 1815)

(=Polymastia robusta)

Range unknown,

found on Georges

Bank, in the Gulf of

Maine and southern

New England

Volume of 40 cm3 Globular with thick base,

body is soft

Most common on upward

facing rock or boulder tops, as

deep as 2300 m (in Europe)

Suberites ficus

(Johnston, 1842)

Arctic south to

Rhode Island,

possibly to Virginia

10‐40 cm diameter

Variable, lobed or globular

cushion, rolls over bottom if

it outgrows its substrate

Attaches to rocks and to small

stones, empty shells, in sandy

or muddy bottom, from 15 to

200 m

(Hecker et al. 1983). This species was also

common in Baltimore Canyon. Brooke et al.

(2017) reported that sponges were commonly

observed on hard substrata in Norfolk and

Baltimore canyons, and that their species

richness exceeded that of corals, with more than

30 different morphotypes of hexactinellid and

demosponges.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Sponges are often a major component of the

epibenthic organisms found on the hard

substrates that dominate seamounts (Samadi et

al. 2007). Cho (2008) reported that the phylum

Porifera represented the most abundant

megabenthic fauna in the New England

Seamount chain (surveys did not include the

four seamounts in U.S. waters). As noted

previously, (Quattrini et al. 2015) observed

numerous and diverse hexactinellid sponges

and demosponges on Mytilus Seamount. Moore

et al. (2003) reported six taxa of sponges on Bear

Seamount, including glass sponges and the

demosponge Geodia sp. The latter forms major

sponge grounds elsewhere in the North

Atlantic, including along the continental slopes

of the Grand Bank and Flemish Cap and

northward along the Labrador Slope to the

southern Davis Strait off Canada (Murillo et al.

2012; Knudby et al. 2013; Kenchington et al.

2014), but high density aggregations have not

been reported in the U.S. Northeast.

II.3. Species Associations with

Deep-Sea Corals and Sponges

Packer et al. (2007) summarized information on

species associations with deep‐sea coral

habitats off the Northeast U.S. While there were

numerous examples of invertebrate associations

with deep‐sea corals in the region, the role of

deep‐sea corals as habitat for commercially

important fish species was unclear.

Recent studies in this region highlight

relationships of invertebrate symbionts and

their octocoral hosts at deep‐sea coral habitats

on the seamounts (Watling et al. 2011). In an

extreme case of host fidelity, Mosher and

Watling (2009) showed that the ophiuroid

Ophiocreas oedipus was found only on the

chrysogorgiid Metallogorgia melanotrichos. The

young brittle stars settle on young corals and

the two species then remain together. The

brittle star may receive some refuge and

feeding benefits from the coral, but the coral’s

relationship to the brittle star appears to be

neutral (Mosher and Watling 2009). Within the

EEZ, these two species were collected from Bear

Seamount at 1491 and 1559 m. Another

ophiuroid, Asteroschema clavigerum, has a close

relationship with Paramuricea sp. and Paragorgia

sp. on both the seamounts and the continental

slope (Cho and Shank 2010). The shrimp

Bathypalaemonella serratipalma as well as the egg

cases of an unknown octopus were found on

Chrysogorgia tricaulis on the seamounts (Pante

and Watling 2011). Additionally, older colonies

of Acanella arbuscula collected from the

seamounts were host to a scale worm (Watling

et al. 2011). De Clippele et al. (2015) reported on

associations of invertebrate fauna with three

gorgonian species (Paragorgia arborea, Primnoa

resedaeformis, and Paramuricea placomus) in

Norway, which are also dominant coral species

in the U.S. Northeast. All hosted numerous

associates, with P. arborea colonies having the

most associated taxa and numbers of

individuals. Cho (2008) found that corals were

the dominant living substrate for other

macroinvertebrates on the New England

Seamount chain; sponges were the second most

important host. See Watling et al. (2011) for

reviews and lists of known invertebrate

symbionts and their octocoral hosts worldwide.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Quattrini et al. (2015) noted that the presence of

certain deep‐sea coral species may influence

crustacean assemblage patterns. For example,

the squat lobster Uroptychus sp. was only

observed on the black coral Parantipathes sp. In

contrast, Munidopsis spp. (squat lobsters in a

different superfamily) utilized a variety of

different coral species as habitat, particularly

those with structurally complex morphologies.

Many other observations suggesting possible

species associations between deep‐sea corals

and other invertebrates are recorded in the

various dive logs and summaries from the

recent Okeanos Explorer surveys.

In all areas explored in the Gulf of Maine,

sponges (e.g., in the genera Polymastia, Iophon,

Phakellia/Axinella) and anemones (e.g.;

Metridium, Urticina) often occurred in high‐

density patches amongst the more extensive

corals on walls and on steep features without

corals (Fig. 8a‐d) (Auster et al. 2014).

Crustaceans such as shrimp, amphipods,

aggregations of krill (Meganyctiphanes

norvegica), and the king crab (Lithodes maja; Fig.

8d) were commonly associated with coral

communities along steep walls and were seen

foraging amongst structure‐forming organisms,

including corals, on the seafloor.

A number of commercially important fish and

shellfish species were observed at the Gulf of

Maine coral sites, including Acadian redfish

(Sebastes fasciatus; juveniles, adults, and

pregnant females; Fig. 7b), haddock

(Melanogrammus aeglefinus; Fig. 7e), pollock

(Pollachius virens), cusk (Brosme brosme),

monkfish (Lophius americanus), Atlantic cod

(Gadus morhua; juveniles, adults; Fig. 7d), silver

hake (Merluccius bilinearis), Atlantic herring

(Clupea harengus), spiny dogfish (Squalus

acanthias), squid (Illex illecebrosus), and lobster

(Homarus americanus). The fish were observed

searching for and catching prey that were also

found among the coral, including shrimp,

amphipods, krill, and other small fish (Auster

et al. 2014). The corals also appeared to provide

refuge from the strong, tidally‐generated

bottom currents.

A cause and effect relationship between

coral/sponge presence and fish populations is

difficult to determine. Our understanding of

relationships between deep‐sea corals and

fishes is both situational and inferential (e.g.,

Baker et al. 2012). This may be particularly true

with regard to the seamount habitats (Auster

2007), although it has been shown, for example,

that false boarfish, Neocyttus helgae, were

associated with basalt habitats colonized by

gorgonian corals and sponges (on both nearly

horizontal basalt sheets and steep cliffs) on Bear

and other seamounts (Moore et al. 2008).

Aggregates of dead coral debris on seamounts

also served as habitat for juveniles of deep‐sea

fish, but observations have been limited (Moore

and Auster 2009). In Norway, Foley et al. (2010)

applied the production‐function approach to

estimate the link between Lophelia pertusa deep‐

sea coral reef habitat and redfish (Sebastes spp.).

Both the carrying capacity and growth rate of

the redfish were found to be functions of deep‐

sea coral habitat. Thus, the authors concluded

that deep‐sea coral reef habitat can be

considered as essential fish habitat. They also

estimated a facultative relationship between

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

deep‐sea coral and Sebastes spp. stocks. Deep‐

sea coral reef (bioherm) habitats do not occur in

the U.S. Northeast, and comparable

quantitative studies on gorgonian‐dominated

habitats have not been carried out.

Recent surveys in the region addressed the

functional role deep‐sea corals play in fish life

history and ecology. Demersal fish distributions

and habitat associations, including the

influence of deep‐sea corals and sponges, were

assessed for Baltimore and Norfolk canyons

(Ross et al. 2015). Although deep‐sea coral and

sponge presence did not significantly influence

fish assemblages in the two canyons, deep‐sea

coral and sponges did increase habitat

complexity, which is an important factor in

determining habitat use for deep‐sea fishes

(Ross et al. 2015). Some of the fishes were

closely associated with the corals, including

Dysommina rugosa (cutthroat eel), Benthocometes

robustus (robust cusk‐eel), and Brosme brosme

(cusk) (Ross et al. 2015). Ross et al. (2015) also

noted unidentified shark egg cases were

frequently entangled in octocorals, especially

the bubblegum coral Paragorgia arborea, while

eggs of snailfish (Liparidae) were also found in

Anthothela sp. and Acanthogorgia sp. Quattrini et

al. (2015) did find that deep‐sea coral species

richness was an important variable in

explaining demersal fish assemblage structure.

These authors hypothesized that corals may

increase fish diversity because the fish use the

corals as habitat, or corals may increase food

resources for fishes. However, Auster (2005,

2007 and references therein) also hypothesizes

that some fish co‐occur with deep‐sea corals

because both taxa do better in areas of

enhanced flows that advect food, and not

because the fish need the corals per se.

Baillon et al. (2012) reported that several species

of sea pens, including Anthoptilum grandiflorum,

Pennatula aculeata, P. grandis, and Halipteris

finmarchica from the Laurentian Channel and

southern Grand Banks in Canada were utilized

as shelter by fish larvae, mainly by those of

redfish (Sebastes spp.). Anthoptilum grandiflorum

may have been of particular importance to

redfish larvae in that study. Both redfish and

these species of sea pens also co‐occur in the

U.S. Northeast region (e.g., Fig. 8f). These

discoveries could have significant research and

management implications for sea pens and

redfish in the Northeast.

The Baillon et al. (2012) study collected sea pens

as trawl bycatch during their routine

multispecies research surveys. A relatively

small number of P. aculeata collected via ROV

from different sites in the U.S. Gulf of Maine in

July/August 2014 yielded no fish larvae. In

2015, 186 individual P. aculeata were collected

as bycatch from the 2015 NMFS Gulf of Maine

northern shrimp survey and subsequently

examined in the laboratory. Redfish larvae were

found on P. aculeata at four stations, either

adhering to the exterior of the colony, or

entrapped within the arms or polyps (Dean et

al. 2016). However, the source of the sea pen

samples for both the Baillon et al. (2012) and

this study was from trawl survey bycatch.

Thus, the possibility that fish larvae were

actually extruded in the net by viviparous ripe

and running redfish, or when the catch was

emptied on the deck, and the larvae then

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

subsequently adhered to the sea pens must be

explored. In June 2016, some stations in the

Gulf of Maine where a positive association had

been found between redfish larvae and P.

aculeata were resampled. This time a small

beam trawl was used as the sampling gear,

increasing the probability that only sea pens,

and not adult redfish, would be captured, thus

eliminating the potential cross contamination

described above. Over 1400 sea pens were

collected over two days of beam trawling over

soft bottoms at 150‐180 m depth. No larval

redfish were found associated with the sea

pens. However, ~80 to 85% of the sea pens

collected were quite small, < 25‐50 mm total

length (adults are upwards of 200‐250 mm).

This suggests a recent recruitment event

occurred, and therefore colonies collected were

probably too small to serve as possible nursery

habitat for larval redfish. Very few larger sea

pens were captured, and those that were caught

were generally tangled in the chain rather than

caught in the net. The beam trawl likely did not

dig deep enough into the sediment to dislodge

these larger animals. Thus, the role of P. aculeata

as possible nursery habitat for larval redfish in

the Gulf of Maine remains uncertain.

II.4. Other new information from

published studies since 2007

Other new findings from the region include

information on coral growth and life history.

For example, Metallogorgia melanotrichos

colonies change form as they grow (Mosher and

Watling 2009). Young colonies have lateral

branches coming off the central axis, while

older colonies lose these branches as the colony

senesces, leaving only a dense crown of

branches.

Life history studies (e.g., reproductive

strategies) on several species of corals found on

both the eastern Canadian shelf and slope

(Newfoundland, Labrador) and off the

Northeastern U.S. (e.g., Acanella, Anthomastus;

see Watling et al. 2011 for review) were also

conducted. The bamboo coral, Acanella

arbuscula, which is abundant in canyons and

slope areas of the Northeast U.S., has separate

sexes (gonochoric) and is likely a broadcast

spawner (Beazley and Kenchington 2012).

Colonies reached sexual maturity at a small size

(~3 cm, corresponding to an age of 3 years) and

larger colonies appeared to be relatively fecund

compared to other gorgonians (Beazley and

Kenchington 2012). The authors nevertheless

concluded that its longevity and the uncertainty

surrounding its dispersal and recruitment,

combined with its vulnerability to bottom‐

contact fishing gear, warranted considering A.

arbuscula as an indicator species or vulnerable

marine ecosystems.

Lawler et al. (2016) reported on the microbial

assemblages of two species of the octocoral

Anthothela from Norfolk and Baltimore

canyons, and Kellogg et al. (2016) examined

Paramuricea placomus bacterial community

diversity in Baltimore Canyon, providing

insights into the functional roles that these

bacteria may play in corals.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

II.5. Research Priorities

Packer et al. (2007) identified a number of deep‐

sea coral research priorities for the region. In

2011, NOAA convened a group of federal, state,

academic, and non‐governmental organization

scientists to identify exploration and research

priorities for deep‐sea coral and sponge

ecosystems in the Northeast region (NOAA

2012b). Key priorities identified, which

continue to be our major priorities going

forward, included the following:

 Locate and characterize deep‐sea coral and

sponge communities:

o Collect and analyze existing data, and

evaluate data gaps

o Develop and refine predictive models for

coral and sponge distribution

o Conduct multibeam sonar surveys to

improve bathymetry data to guide field

research and for use in spatially‐explicit

models (canyons and deepwater areas of

the Gulf of Maine)

o Conduct site‐specific research designed to

locate and characterize deep‐sea coral and

sponge habitats

 Understand the biology, biodiversity and

ecology of deep‐sea corals and sponges,

including:

o Species identification

o Population connectivity

o Age and growth rates

o Functional role

 Understand natural and human impacts on

deep‐sea coral and sponge ecosystems:

o Disturbance effects and recovery

o Economic and social values associated

with these coral and sponge

communities

New research conducted since 2011 has

addressed a number of these priorities for deep‐

sea corals, resulting in extensive multibeam

mapping of the shelf‐edge and slope, the first

comprehensive predictive habitat models for

deep‐sea corals, extensive new exploratory

surveys, and studies of community ecology

(trophic and population structure) and coral

biology (age, growth, life histories). Studies of

various aspects of the biology and ecology of

key species (e.g., Primnoa resedaeformis) are

underway. However, much additional work

remains to be done. Studies on deep‐sea

sponges, including taxonomy, distribution, life

history, and habitat functions, are almost totally

lacking.

III. Update on Management

of Deep-Sea Corals and

Sponges

III.1. New Information on Impacts

and Stressors

III.1.i – Fishing

Fishing impacts to deep‐sea corals in the

Northeast region were discussed previously

(Packer et al. 2007), but this is an active field of

research, and new field and modeling studies

have been published since 2007. Much of this

work is applicable to the Northeast region and

informs interpretation of observations of

obvious gear impacts to regional deep‐sea coral

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

and sponge habitats. In any case, the

conclusions about the nature of these impacts

are generally similar to earlier work. Because

the predominate industrial use of the deep‐sea

in the Northeast is fishing, impacts from

bottom‐contact fishing are still considered the

greatest and most immediate threat to deep‐sea

corals and sponges in this region. The

seriousness of the threat is reflected in the

proactive work of the Fishery Management

Councils in pursuing the current management

options, discussed below.

Most bottom‐contact fisheries in the Northeast

U.S. are conducted on the continental shelf,

including the Gulf of Maine and Georges Bank,

with some fishing effort around the shelf/slope

break. Considering the major areas of coral

distribution described above, the Gulf of Maine

and the upper portion of canyons and slope are

subject to the highest fishing pressures. Fishing

for monkfish (largely by trawl, with some

gillnet fishing) and tilefish (predominantly by

bottom longlines) occurs predominantly on the

shelf, but can extend into deeper waters, as can

the squid, whiting, and butterfish trawl

fisheries. Lobster pots are used to harvest

American lobster, as well as Jonah crab, along

the shelf break. A small deep‐sea red crab

(Chaceon quinquedens) fishery is active in deeper

waters, deploying strings of 90‐120 pots

primarily at depths from 400‐800 m (NEFMC

2002), targeting approximately the 640 m depth

contour.

During the 2012‐2015 Gulf of Maine exploratory

surveys, areas exhibiting direct impacts from

fishing activities were observed at sites in

Western and Central Jordan Basin, Outer

Schoodic Ridge, and Georges Basin (Auster et

al. 2014). In steep areas, paths or tracks,

consistent with the setting or recovery of trap

gear, were denuded of corals and associated

fauna (Fig. 9a). The peaks of some ridges and

nearly horizontal sections of wider rock

outcrops were also denuded. Tracks observed

here were consistent with impacts from mobile

fishing gear (Fig. 9b,c). Some coral patches

exhibited damage in the form of live colonies

with disjunct size class structure, suggesting

past impacts (Fig. 9d). In areas such as Georges

Basin, colonies of Paramuricea placomus and

associated species were often small and

virtually all occurred in physical refuges such

as cracks and crevices of outcrops and along the

sediment‐rock interface of large cobbles and

boulders. Of note is that the sea star Hippasteria

phrygiana was observed eating or preying on P.

resedaeformis colonies at the Outer Schoodic

Ridge site. These sea stars were seen on living

coral colonies that had been detached from rock

walls and were laying on the seafloor, possibly

due to fishing activity, as one was seen next to

an abandoned fishing net. Opportunistic

predation by H. phrygiana has also been noted

in Alaska on Primnoa pacifica that had been

injured or detached by fishing gear (Stone et al.

2015). This may indicate that coral damaged by

fishing gear interactions are at an increased risk

of predation by sea stars, thus further reducing

the chances that a coral colony will recover

from gear‐related injuries and impacts.

Knight et al. (2006) analyzed recovery of

epibenthic megafauna in predominantly soft‐

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Figure 9. Examples of fishing gear impacts to coral habitats in the Gulf of Maine (Auster et al. 2014 and

unpublished images). (a) Denuded bottom (sediment covered hard bottom) in Western Jordan Basin (114

Bump) consistent with fixed gear impacts. (b) Denuded ridge in Central Jordan Basin consistent with mobile

gear impacts. (c) Denuded ridge in Georges Basin consistent with mobile gear impacts. (d) Example of oddly

shaped corals with disjointed size class structure, possibly due to damage, in Western Jordan Basin.

sediment shelf areas in the Western Gulf of

Maine that had been closed to fishing for 6

years. The closed sites showed higher average

taxonomic abundance than fished sites – with

unidentified sponges and a tunicate (Mogula

sp.) responsible for approximately 83% of the

difference. This suggests that sponges are

susceptible to impacts from bottom‐contact

fishing, and at least certain species may recover

more quickly than corals.

The annual number of interactions between

fishing gear and deep‐sea corals and sponges is

not known, but bycatch data indicate that a

relatively small number of trips interact with

deep‐sea corals. In 2011 NOAA’s National

Marine Fisheries Service (NMFS) granted the

Maine Department of Marine Resources an

exempted fishing permit for redfish in order to

conduct a baseline catch and bycatch evaluation

in and around Wilkinson Basin and elsewhere

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

in the central Gulf of Maine. Many smaller

individuals escape from the 6.5 inch mesh nets

required by groundfish regulations. The

experimental fishing used nets with smaller, 4.5

inch mesh liners in the cod end and targeted

schools of redfish that congregate on ʺbumpsʺ 

or pinnacles that occur in the normally deep,

muddy areas in the central Gulf of Maine. Since

redfish seek shelter near structure‐forming

organisms such as deep‐sea corals and sponges,

as well as boulder reefs (Packer et al. 2007),

concerns were raised by NMFS that the smaller

mesh nets would increase the probability of

bycatch of deep‐sea corals. NMFS determined

that the project could have an adverse effect on

EFH, particularly on any deep‐sea corals found

there. Therefore, they requested that deep‐sea

coral bycatch be carefully monitored to enhance

the understanding of deep‐sea coral

distribution in the Gulf of Maine and the

potential effects of an expanded redfish fishery

on deep‐sea corals. However, by the end of the

project the only coral by‐catch was that of a

single specimen of the common sea pen,

Pennatula aculeata, which, as stated previously,

is ubiquitous in muddy areas of the Gulf of

Maine. It seems that none of the project tows

overlapped with any known soft coral habitats,

even though the current redfish fishery appears

to operate in the vicinity of these habitats.

Several years ago, NMFS Northeast Fisheries

Science Center’s (NEFSC) fishery independent

surveys and Northeast Fisheries Observer

Program (NEFOP) were assessed for potential

coral bycatch. As discussed previously (Packer

et al. 2007), the spring/fall NEFSC groundfish

trawl and scallop dredge surveys conducted in

the region do not “catch” deep corals in any

meaningful quantities, nor is any deep coral

that may be brought on deck recorded in a

significant quantitative way. The NEFSC

groundfish trawl surveys loosely describe and

roughly quantify any substrate (rock, shell, etc.)

or non‐coded invertebrate species. Although

this bycatch information could possibly be

useful as presence/absence data, since deep‐sea

corals are not the focus of the bottom trawl

surveys, these data should of course be used

with caution (John Galbraith, NMFS, NEFSC,

Woods Hole Laboratory, Woods Hole, MA,

pers. comm.).

Outside the Gulf of Maine, the general lack of

deep‐sea coral in both the groundfish trawl and

scallop dredge surveys may be a function of the

surveys fishing in waters shallower than where

the larger deep‐sea coral species are likely to

occur; i.e., the groundfish survey fishes from

about 9‐366 m depth, while nearly all the

scallop surveys fish < 100 m and all are < 140 m.

In some areas in the Gulf of Maine and Georges

Bank, these larger corals (e.g., Paragorgia

arborea, Primnoa resedaeformis) may have already

been “fished out” in the survey areas during

the 19th and 20th centuries (Packer et al. 2007).

In Canadian waters near the Northeast

Channel, but within the survey region, there is

a deep‐sea coral protection area that is closed to

fishing. J. Galbraith (NEFSC, pers. comm.)

stated that this was the only area he could

remember where any amount of coral was

encountered. Anecdotal accounts from the time

period before the groundfish survey began

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

(1950s or early 1960s) reference an area on

Georges Bank called ʺThe Treesʺ where large

corals existed in shallower water before being

eventually cleared out, supposedly by foreign

trawling vessels. There are also anecdotal

accounts from Gloucester fishermen who said

that they used bottom gear on Georges Bank to

first knock down the “trees” before fishing for

large adult redfish.

The fishery dependent deep‐sea coral bycatch

data collected from the NEFOP used to suffer

many of the same problems as the groundfish

trawl and scallop dredge surveys. First, it

should be noted that only 10‐40% of all fishing

effort is observed by “observers” onboard

vessels, depending on the fishery, and a grand

average may be somewhere around 10%. A

small NEFOP database of coral bycatch

collected from 1994‐2009 showed only 39

confirmed coral entries. Two of these entries

were labeled ‘Astrangia’ (a solitary stony cup

coral) and 10 additional entries were labeled

stony corals. Basic information about the haul

(gear type, year, month, depth, and geographic

coordinates) was included. Gear used included

otter trawls, scallop dredges, and gill nets, at

depths from 5.5‐253 m (depths were taken at

the beginning of a tow or set). Estimated or

actual weights for the coral in a given haul

ranged from 0.05 − 22.7 kg. No specimens or

photographs were included.

In 2013, the NEFOP training curriculum and

associated sampling protocols were changed to

improve deep‐sea bycatch identification,

retention, enumeration, and documentation

(Lewandowski et al. 2016). This included the

use of a Northeast deep‐sea coral identification

guide for the onboard observers, and

standardized recording, sampling, and

preservation procedures. Since implementation

of the new protocols, although deep‐sea coral

bycatch is still low, the number of recorded and

verified samples has increased, and

photographic records and samples are being

stored via the NEFOP Species Verification

Program (Lewandowski et al. 2016). These were

recently reviewed and classified by Northeast

deep‐sea coral experts. Several structure‐

forming gorgonians, as well as sea pens, were

documented in bycatch by at‐sea observers.

Improved NEFOP fishery dependent deep‐sea

coral bycatch data will lead to a better

understanding of fisheries and deep‐sea coral

interactions and impacts, and guide

conservation efforts of deep‐sea corals habitats

in the Northeast. Of course, bycatch only

reveals what is not lost below the net, or not

lost through the mesh, or not destroyed within

the net (see Auster et al. 2011 on this issue).

III.1.ii – Other stressors

Oil and gas drilling, while currently not

conducted within the region, took on an added

urgency with the lifting of the ban on oil

drilling off Virginia in 2010; hence, the BOEM‐

funded study discussed above. The Mid‐

Atlantic was removed from the 2017‐2022 Outer

Continental Shelf Oil and Gas Leasing

Proposed Program (BOEM 2016), but this

decision could be revisited. On June 29 2017,

the Secretary of the Interior announced a

review of the 2017‐2022 Program to include all

26 planning areas. Ocean acidification, although

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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not discussed in Packer et al. (2007), is

obviously a major threat to all corals worldwide

(e.g., for tropical coral reefs, see Kleypas and

Yates 2009). Research on ocean acidification in

relation to deep‐sea corals has begun (e.g.,

Form and Riebesell 2012; Lunden et al. 2014),

but has focused primarily on stony corals (i.e.,

Lophelia pertusa). There have been no studies on

the dominant coral species in the Northeast

region.

Other activities that impact the seafloor and

pose potential threats to deep‐sea corals and

sponge habitats in the region include

deployment of seafloor cables and offshore

renewable energy installations. New York is a

major hub for underwater communication

cables, but there is little information on impacts

to biogenic habitats. Potential offshore wind

development has become a major driver for

more comprehensive ocean planning within the

region. The Bureau of Ocean Energy

management (BOEM) has awarded leases in

specific areas from Massachusetts to Virginia.

In 2016, the Block Island Wind Farm off Rhode

Island began operations as the nation’s first

offshore wind installation. This and future

wind development is likely to be conducted on

the continental shelf relatively close to shore,

and thus less likely to impact major deep‐sea

coral and sponge habitats, although it could

affect shallower water sponge populations.

Marine debris and trash are also a threat to

corals in this region, and have been noted in

some of the recent surveys. Quattrini et al.

(2015) found an estimated 0.002‐0.130 items/10

m2, including derelict fishing gear (ghost traps,

lines, hooks) and trash (cans, bottles, plastic

bags, etc.). The highest numbers came from one

of the minor, unnamed canyons. At least 12

deep‐sea coral colonies were tangled in debris,

including five gorgonian colonies (Paramuricea

spp., Thouarella grasshoffi) that suffered varying

degrees of injury or death. Brooke et al. (2017)

noted marine debris on almost every dive in

Baltimore and Norfolk canyons, often near the

canyon heads on rock outcrops; canyon heads

are where most fishing activity would take

place. During the Gulf of Maine exploratory

surveys, little debris was seen, which was

surprising considering the number of vessels

traversing the survey areas. Debris observed

included lines, nets, trash, and in one instance

in Western Jordan Basin, a large piece of plastic

sheeting that covered a boulder with several

large colonies of Paramuricea placomus.

Packer et al. (2007) discussed potential impacts

of an invasive colonial tunicate (Didemnum sp.

A) found on gravel and cobble substrates on

Georges Bank. This species has since been

identified as Didemnum vexillum (Lengyel 2009).

The progress of this species is being monitored,

as it can have a major effect on the benthic

invertebrate communities of Georges Bank

(Kaplan et al. 2017a, b), but it has not been

reported in the primary habitats of high‐density

deep‐sea coral or sponge communities (e.g.,

Bullard et al. 2007).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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III.2. New or Planned

Management Actions

III.2.i – Northeast Canyons and Seamounts

Marine National Monument

On September 15, 2016, President Obama

designated the first marine national monument

in the Atlantic Ocean, the Northeast Canyons

and Seamounts Marine National Monument,

using his authority under the Antiquities Act of

1906. The monument consists of two units (Fig.

10), representing distinct geological features

that support vulnerable ecological

communities, including deep‐sea coral and

sponge communities. The Canyons Unit covers

approximately 2,437 km2 (941 square miles) on

the edge of Georges Bank, including

Oceanographer, Gilbert, and Lydonia

submarine canyons. The Seamounts Unit

encompasses 10,287 km2 (3,972 square miles)

and includes the four New England Seamounts

in the U.S. EEZ: Bear, Mytilus, Physalia, and

Retriever. The presidential proclamation

specifically referenced deep‐sea corals, along

with “other structure‐forming fauna such as

sponges and anemones,” as resources that

“create a foundation for vibrant deep‐sea

ecosystems” and are extremely sensitive to

disturbance from extractive activities.

The new monument will be managed jointly by

the Department of the Interior and the

Department of Commerce (through NOAA).

The proclamation creating the Monument

identifies a number of prohibitions designed to

help protect monument ecosystems, including

prohibiting oil, gas, and mineral exploration

and development. Commercial fishing in the

monument is prohibited with the exception of

existing red crab and lobster fisheries, which

were granted a grace period for up to seven

years. Other activities, such as cable

deployment and maintenance, scientific

activities, and recreational fishing will be

permitted as long as they are determined to be

consistent with the conservation goals of the

Monument. The Department of the Interior and

NOAA will jointly develop a management plan

for the Monument within three years. Recent

marine national monument designations are

undergoing review by the Department of the

Interior and NOAA, in response to an April 26,

2017 Executive Order.

III.2.ii – Fisheries management

In the Northeast region, fisheries in Federal

waters are primarily managed by NOAA’s

National Marine Fisheries Service (NMFS)

under fishery management plans (FMPs)

developed by the New England and Mid‐

Atlantic Fishery Management Councils

(NEFMC, MAFMC). Recent work by both

regional Councils has been facilitated by the

new exploratory surveys and data on the

distribution of deep‐sea corals. As described

above, most of these data were collected

between 2012 and 2015. The Councils, in

collaboration with NMFS, recommended

priority targets to ensure that results would be

relevant to managers. Additionally, in 2011,

NMFS, in collaboration with the two regional

Councils, initiated a partnership to develop and

implement a strategy for deep‐sea coral

conservation as part of NOAA’s Habitat

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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Figure 10. Deep‐sea coral protection zones for the Northeast region enacted in the Mid‐Atlantic and being

considered for New England. Broad zones are shaded; discrete zones on seamounts, in canyons, and in the

Gulf of Maine, are outlined. Alvin Canyon straddles the New England/Mid‐Atlantic inter‐council boundary.

New England zone boundaries will likely change as the amendment is finalized based on stakeholder feedback.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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Blueprint program. The goal was to ensure

effective, long‐term conservation of deep‐sea

coral habitats with existing management and

scientific resources as well as provide a

template to integrate habitat conservation and

science into the Council process. This program

took advantage of deep‐sea coral science and

management activities that were already

underway in the region, including the 2013‐

2015 NOAA Northeast Deep‐sea Coral and

Sponge Initiative. Another objective of this

program was to implement a coral conservation

strategy with federal offshore energy licensing

agencies (e.g., BOEM) using existing

coordination and consultation procedures.

There are two primary authorities in the

Magnuson Stevens Fishery Conservation and

Management Act (MSA) that can be used to

protect deep‐sea coral and/or sponge habitats.

Under the Essential Fish Habitat (EFH)

authority, fishing restrictions should be enacted

to minimize, to the extent practicable, the

adverse effects of fishing on corals or sponges

that are considered a component of EFH for a

managed fisheries species (MSA section 305(b)).

Actions taken under the EFH authority

generally occur within areas that are designated

as EFH. The NEFMC used this authority in 2005

to protect deep‐sea corals and associated

habitat features in Lydonia and Oceanographer

canyons from fishing activity occurring under a

monkfish “days‐at‐sea” permit. The MAFMC

used the same authority in 2008 to close these

two canyons to bottom trawling by vessels in

the squid, mackerel, and butterfish fisheries via

Amendment 9 to that FMP. In 2009 the

MAFMC closed Lydonia, Oceanographer,

Veatch, and Norfolk Canyons (as tilefish Gear

Restricted Areas) to mobile bottom tending

gear through Amendment 1 to the tilefish FMP,

with the intention of protecting vulnerable

tilefish habitat (clay outcroppings also known

as “pueblo” habitats).

Areas outside designated EFH may also be

managed under the EFH authority, but the

scope of this authority is presumably not

limitless. In the Northeast region, coral

distributions extend into waters deeper than

the bounds of managed species and their

designated EFH. The deep‐sea coral

discretionary provisions (MSA Section 303(b))

from the 2007 MSA reauthorization provide an

additional and more flexible mechanism by

which to protect deep‐sea corals from physical

damage by fishing gear. This discretionary

authority allows Councils to designate

protection zones anywhere corals have been

identified by the Deep Sea Coral Research and

Technology Program and implement measures

to protect corals within those zones, provided

that long term sustainable use of fishery

resources has been considered, and there is a

nexus to Council‐managed fisheries (NOAA

2010; Sutter 2014). This authority has become

the primary basis for a range of coral protection

measures enacted by the MAFMC and

currently under development by the NEFMC

(Fig. 10). The measures define the boundaries of

coral protection zones and implement fishing

restrictions within them. In addition to the EFH

and discretionary authorities, National

Standard 9 of the MSA requires federal fishery

management plans to minimize bycatch to the

extent practicable.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

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Substantial spatial overlap exists between

continental slope fisheries managed by the

NEFMC and MAFMC. This overlap affects the

range of coral alternatives developed and their

potential impacts on fishing activities. The

NEFMC began development of a deep‐sea coral

amendment in 2010 that included coral areas

from the Gulf of Maine to Norfolk Canyon, off

Virginia. In 2012, the MAFMC decided to

initiate their own FMP amendment to protect

deep‐sea corals in the Mid‐Atlantic region

(roughly south of Alvin Canyon). These two

Councils, along with the South Atlantic Fishery

Management Council, then drafted a

memorandum of understanding (MOU) to

coordinate deep‐sea coral management

activities. Also during 2012, the NEFMC

decided to develop their deep‐sea coral

management measures in a standalone coral‐

focused amendment, independent from a much

more comprehensive omnibus EFH

amendment.

In 2015, the MAFMC recommended specific

proposals to NMFS to prohibit the use of most

types of bottom‐tending fishing gear within a

99,000 km2 (~38,000 square miles) area on the

outer continental shelf, slope, and canyons to

the outer boundary of the EEZ. This was

approved by NOAA in 2016, marking the first

use of the MSA deep‐sea coral discretionary

authority. The new management zones are

officially known as the “Frank R. Lautenberg

Deep‐Sea Coral Protection Area.” Senator

Lautenberg, a senator from New Jersey, was

responsible for, and authored, several

important pieces of ocean conservation

legislation, including several provisions in the

reauthorized MSA, including the discretionary

provisions described above.

The Frank R. Lautenberg Deep‐Sea Coral

Protection Area includes two types of zones.

‘Discrete’ zones protect defined areas of

canyons and canyon complexes based on the

best available information on known coral

distributions or outputs of predictive models

that rank the likely presence of suitable coral

habitats. A precautionary ‘broad’ zone protects

a large area of deepwater habitats extending

from approximately 450 m on the slope to the

outer limits of the U.S. EEZ (Fig. 10). The

objective is to protect corals by limiting future

expansion of bottom fishing in an area that is

largely outside the footprint of current fishing

activity. Both zones restrict most bottom‐

tending gears, but the red crab fishery is

exempted for a period of at least two years in

the discrete zones and indefinitely in the broad

zones, and the action does not restrict

lobster/Jonah crab traps in either type of coral

zone. The lobster fishery is primarily managed

by the Atlantic States Marine Fisheries

Commission. This dual discrete/broad

framework follows the approach of NOAA’s

Strategic Plan for Deep‐Sea Coral and Sponge

Ecosystems (NOAA 2010), and provides

flexibility to revise fishing restrictions

independently in the various areas.

The NEFMC is still developing and evaluating

management approaches and will likely make

final recommendations to NMFS during late

2017 or early 2018. The council has taken a

similar approach to that of the MAFMC,

including a precautionary ‘broad’ zone in

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

deeper water. Discrete area options that are

being considered in New England include

seven areas in the Gulf of Maine, a number of

submarine canyons and canyon complexes, and

the four New England Seamounts in the U.S.

EEZ southeast of Georges Bank.

III.2.iii – Other Resource Management and

Ocean Policy Efforts

The Mid‐Atlantic Regional Council on the

Ocean (MARCO) was created by the Governors

of five Mid‐Atlantic States (NY, NJ, DE, MD,

VA) to improve regional coordination of shared

ocean issues and to address the ocean

environment across the states as a whole

ecosystem, through the principles of ecosystem‐

based management (Capobianco 2011). The first

MARCO priority is to coordinate the protection

of important habitats and sensitive and unique

offshore areas, such as the submarine canyons

and deep‐sea coral and sponge habitats, on a

regional scale. To facilitate habitat protection

actions by its partners, MARCO developed an

online portal (http://midatlanticocean.org/data‐

portal/) that displays geospatial information

obtained from sources such as NOAA’s deep‐

sea coral and sponge database to aid in

identifying regionally‐important habitats such

as the canyons (Capobianco 2011). The high‐

resolution multibeam bathymetric surveys of

the slope and canyons conducted by NOAA

were done in part to support MARCO’s habitat

mapping and protection goals. MARCO has

been following the work of the two Councils in

protecting submarine canyon habitats, and

facilitated the engagement of multiple federal

entities and stakeholders with an interest in

these habitats to ensure that all existing

resources and authorities were leveraged.

Similar planning and geospatial data gathering

efforts are underway in New England as well,

and coral‐related data products including

habitat suitability model outputs are available

through the Northeast Ocean Data Portal

(http://www.northeastoceandata.org). In

December, 2016, the National Ocean Council

certified the Northeast and Mid‐Atlantic Ocean

Plans.

NOAA’s National Marine Sanctuary Program

also received nominations from communities

for new National Marine Sanctuaries for

Baltimore Canyon and Hudson Canyon

(http://www.nominate.noaa.gov/nominations/).

In both cases, deep‐sea coral resources were

identified in the justification for consideration

for sanctuary status. The Baltimore Canyon

proposal was withdrawn in January 2017, but

the Hudson proposal was moved into

candidate status at NOAA during February

2017.

V. Conclusions

Mapping and exploratory surveys since 2010

have covered almost all the submarine canyons

off the Northeast coast as well as areas of the

continental slope, the seamounts within the

EEZ, and significant areas in the Gulf of Maine.

Considering the state of our knowledge of

deep‐sea coral and sponge habitats prior to

these surveys, the results documented herein

have increased our knowledge base of

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

Northeast deep‐sea coral and sponge habitats

exponentially. They have also generated much

interest from the public and media (e.g., see

NRDC 2014 for some 2011‐2014 canyon and

seamount expedition highlights). However,

knowledge gaps still exist for many of these

organisms, especially for the sponges, and some

basic questions about distribution, abundance,

taxonomy, connectivity, life history, population

biology, functional role, effects and scale of

disturbance, ecological resilience, etc. remain

unanswered. Nevertheless, the new surveys

have contributed greatly to our regional

knowledge of deep‐sea coral diversity,

distribution, and habitat characteristics.

From a management perspective, these results

prompted the NEFMC and MAFMC to revisit

their previous lists of potential deep‐sea coral

protections zones, especially those for the

submarine canyons and the Gulf of Maine.

Now that most of these areas have been

surveyed and deep‐sea corals were observed in

almost all of them, these areas can be

confidently classified as “assessed” and as

“suitable” deep‐sea coral habitat. The Councils,

therefore, were then able to recommend

additional areas for consideration as deep‐sea

coral protection zones/ management areas.

Having this information led directly to the

creation of the Frank R. Lautenberg Deep‐Sea

Coral Protection Area in the mid‐Atlantic, will

support the decision‐making process for the

creation of deep‐sea coral protected areas by

the NEFMC, and was one of the major factors

behind the creation of the Northeast Canyons

and Seamounts Marine National Monument. In

addition, a high correlation between recent

coral observations and locations predicted to be

suitable coral habitat has increased confidence

in using the regional deep‐sea coral habitat

suitability model to inform current and future

management decisions of the Councils. Both the

NEFMC and MAFMC have used and continue

to use the information and results gathered

from the recent exploratory surveys, the

historical database, and the habitat suitability

model to draft management alternatives to

designate deep‐sea coral zones in the Northeast

and Mid‐Atlantic and implement fishing

restrictions necessary to protect the deep‐sea

corals within those zones.

V. Acknowledgements

The authors thank those people from other

NOAA Line Offices that were either part of the

Bigelow/Okeanos/Hassler mapping/survey cruises

and/or led and provided assistance/funding for

modeling, surveys, etc. including Mashkoor

Malik, Jeremy Potter, Elizabeth Lobecker,

Nathan Keith, LCDR Ben Evans, LT Sam

Greenway, and Tim Shank from WHOI. The

authors also thank the Gulf of Maine survey

scientists, including Peter Auster, Rhian Waller,

Rich Langton, and Steve Auscavitch; the crew

and engineers of the Connecticut, ISIS2, and

Kraken 2, including Ivar Babb; and the Sandy

Hook folks who help get the data out there:

Donna Johnson, Jeff Pessutti, Barry Shafer,

Christine Shafer, and Caren Martins.

Additional thanks to Tania Lewandowski

(NEFOP). Finally, we thank Rich Langton

(NEFSC), David Stevenson (GARFO), Stephen

Cairns (Smithsonian NMNH), Andrea Quattrini

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE NORTHEAST UNITED STATES

  

(Harvey Mudd College), Sandra Brooke

(Florida State Univ.), Rhian Waller (U. Maine),

Peter Auster (U. Conn.), Kiley Dancy

(MAFMC), and especially Tom Hourigan for

helpful reviews and comments. Figure 5d

courtesy of Source: Deepwater Canyons 2012

Expedition NOAA‐OER/BOEM/USGS. Figures

5e,f,l courtesy of NOAA Okeanos Explorer

Program, 2013 Northeast U.S. Canyons

Expedition. Figures 5g,h,i courtesy of NOAA

Okeanos Explorer Program, Our Deepwater

Backyard: Exploring Atlantic Canyons and

Seamounts. Gulf of Maine Figures 7‐9 courtesy

of NOAA/UConn. – NURTEC/UMaine.

This chapter is dedicated to the memory of

Brian Kinlan, whose presence is deeply missed

but whose ideas and passion live on.

Online Annex – Comprehensive list of deep‐

sea corals in the U.S. Northeast region link:

https://deepseacoraldata.noaa.gov/library/2017‐

state‐of‐deep‐sea‐corals‐report

VI. Literature Cited

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Geographic Distribution 

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Appendix – Deep-Sea Coral and Sponge Distribution Maps 

Map 1. Locations of black corals (Order Antipatharia) recorded in the National Deep‐Sea Coral and

Sponge Database (as of October 2017).

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Map 2. Locations of stony corals (Order Scleractinia) recorded in the National Deep‐Sea Coral and Sponge

Database (as of October 2017).

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Map 3. Locations of soft corals (Order Alcyonacea) recorded in the National Deep‐Sea Coral and Sponge

Database (as of October 2017).

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Map 4. Locations of gorgonians (Order Alcyonacea, in part – formerly Order Gorgonacea) recorded in the

National Deep‐Sea Coral and Sponge Database (as of October 2017).

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Map 5. Locations of sea pens (Order Pennatulacea) recorded in the National Deep‐Sea Coral and Sponge

Database (as of October 2017).

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Map 6. Locations of sponges (Phylum Porifera) recorded in the National Deep‐Sea Coral and Sponge Database

(as of October 2017). Although sponges occur throughout the region, observations have not been

systematically recorded and organized, so records in the database are currently limited.

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Leiopathes black coral colony on the West Florida escarpment in the Gulf of Mexico. Red lasers

visible near the middle of the image indicate that the basal axis of the colony is about 1.7 cm in

diameter. This measurement suggests the coral colony is between 400‐1000 years old, based on

published growth rates for black corals in the Gulf of Mexico (Prouty et. al, 2011). Black corals

in the genus Leiopathes are among the longest‐lived marine organisms, reaching ages of

thousands of years. Courtesy of the NOAA Office of Ocean Exploration and Research.

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AGE, GROWTH RATES, AND PALEOCLIMATE STUDIES IN DEEP-SEA CORALS OF THE U.S.

• • •

AGE, GROWTH 

RATES, AND 

PALEOCLIMATE 

STUDIES IN DEEPSEA CORALS OF 

THE U.S. 

I. Introduction

Deep-water corals are some of the slowest growing, 

longest-lived skeletal accreting marine organisms. 

These habitat-forming species support diverse faunal 

assemblages that include commercially and ecologically 

important organisms. Therefore, effective management 

and conservation strategies for deep-sea corals can be 

informed by precise and accurate age, growth rate, 

and lifespan characteristics for proper assessment of 

vulnerability and recovery from perturbations. This is 

especially true for the small number of commercially 

valuable, and potentially endangered, species that are 

part of the black and precious coral fisheries (Tsounis 

et al. 2010). In addition to evaluating time scales of 

recovery from disturbance or exploitation, accurate age 

and growth estimates are essential for understanding 

the life history and ecology of these habitat-forming 

corals. Given that longevity is a key factor for 

population maintenance and fishery sustainability, 

partly due to limited and complex genetic flow among 

coral populations separated by great distances, accurate 

age structure for these deep-sea coral communities is 

essential for proper, long-term resource management.

The importance of accurate age and growth 

characteristics has another important utility in marine 

sciences. Many deep-sea corals have been useful as 

Nancy G. Prouty1*, 

E. Brendan Roark2,

Allen H. Andrews3,

Laura F. Robinson4,

Tessa Hill5, Owen 

Sherwood6, Branwen 

Williams7, Thomas 

Guilderson8, and 

Stewart Fallon9 

• • •

1

 Pacific Coastal and Marine 

Science Center, U.S. Geological 

Survey, Santa Cruz, CA 

* Corresponding Author:

NProuty@usgs.gov

2 Department of Geography, Texas 

A&M University, College Station

3 NOAA Pacific Islands Fisheries 

Science Center, Honolulu, HI

4 School of Earth Sciences, 

University of Bristol, U.K.

5 Bodega Marine Laboratory, 

University of California, Davis

6 Institute of Arctic and Alpine 

Research, University of Colorado, 

Boulder

7 W.M. Keck Science Department, 

Claremont Colleges, Claremont, 

CA

8 Lawrence Livermore National 

Laboratory; University of 

California, Santa Cruz, CA

9 Research School of Earth 

Sciences, Australian National 

University, Canberra, Australia
AGE, GROWTH RATES, AND PALEOCLIMATE STUDIES IN DEEP-SEA CORALS OF THE U.S.

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300

biogeochemical proxies that provide a unique 

view of marine climate and environmental 

change over time (e.g., Adkins et al. 1998, 

Robinson et al. 2005, LaVigne et al. 2011). 

Similar to trees on land, many corals have 

concentric growth rings that allow scientists 

to track growth patterns, which can be used 

to look back into climate history over the 

coral’s lifespan using stable and radio-isotope 

techniques. Here we summarize recent 

developments in determining age and growth 

characteristics for structure-forming deepsea corals, many of which are vulnerable 

to disturbance, with highlights on recent 

advances in paleoclimate reconstruction 

efforts using deep-sea corals.

II. Age and Growth Rates

Age and growth rates of deep-sea corals 

have been determined by methods that differ 

greatly in approach and experimental design. 

These methods include tagging and outgrowth 

observations (often limited to shallower 

depths; Grigg 1976, Stone and Wing 2001, 

Peck and Brockington 2013), sclerochronology 

(counting of growth rings in the axial skeleton; 

Grigg 1974, Wilson et al. 2002, Marschal et al. 

2004, Tracey et al. 2007, Aranha et al. 2013), 

and by radiometric techniques (Uranium/

Thorium, Lead-210 and radiocarbon (14C) 

dating; Cheng et al. 2000, Adkins et al. 

2002, Andrews et al. 2009, Roark et al. 2009, 

Carreiro-Silva et al. 2013, Aranha et al. 2014). 

When evaluating reported age and growth 

rates, it is important to evaluate the strengths 

and weakness of each method along with 

its applicability to the potential lifespan 

(Figure 1). These may include the limits of 

methodological dating and precision, as well 

as skeletal morphology and mineralogy. For 

example, in traditional tagging studies it is 

difficult to get accurate in-situ measurements

and it can take many years to see measurable

growth (Grigg 1976). Sclerochronology can

provide estimates of age from visible growth

rings in the skeletal structure (Figure 2), but

this approach requires validation of the ring

formation periodicity (Andrews et al. 2002;

Sherwood et al. 2005). Radiometric techniques

have different ranges for effective dating and

usually require assumptions. With recent

advances in Uranium/Thorium dating, it is

possible to determine the age of aragonitic (a

form of calcium carbonate) corals from less

than a few decades old up to 600,000 years in

age (Cheng et al. 2000). Typical uncertainty is

variable and dependent upon various factors,

but recent developments have reduced the

uncertainty (Edwards et al. 1987, McCulloch

and Mortimer 2008). Lead-210 dating is

typically applicable to the last ~100 years with

uncertainties as low as ±10 years and is most

applicable to determining a mean growth

rate for the entire colony (Andrews et al.

2009). The range of conventional radiocarbon

dating is from modern time (defined as 1950

AD) to ~50,000 years BP. While the method is

effective, challenges exist in most applications

because the atmospheric concentration of

radiocarbon has varied over time (Stuiver

and Brazunias 1993, Reimer et al. 2009). In

the marine environment, radiocarbon ages

are also corrected for the difference between

the atmospheric radiocarbon content and

the local radiocarbon content of the surface

ocean, also known as a “reservoir correction.”

Age determination in deep-sea corals using

radiocarbon dating is most applicable to

growth occurring over centuries to millennia

for living specimens, and extends into fossil

specimens in the tens of thousands of years

(Guilderson et al. 2005). A second form of

radiocarbon dating for some living deepsea corals involves the identification of the
301

Figure 2. Ultraviolet light illuminates the growth rings in 

a cross-section of a 44-year-old deep-sea coral (Primnoa 

resedaeformis) collected off the coast of Newfoundland 

at 400 meters. Similar to trees, cross-sections reveal coralgrowth rings (photo by Owen Sherwood).

Figure 1. Maximum lifespan (years) of several key deep-sea corals (blue), and the relative time span of 

efficacy for different dating techniques (red). Typical dating uncertainties listed for dating techniques.

AGE, GROWTH RATES, AND PALEOCLIMATE STUDIES IN DEEP-SEA CORALS OF THE U.S.

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Figure 3. Distribution of bomb-derived radiocarbon (r14C) in proteinaceous deep-sea corals illustrating 

the uptake and delivery of elevated r14C in food sources (e.g., rapidly exported surface derived organic 

matter) to the deep-sea coral community. In comparison, the r14C of the carbonate portion of the bamboo 

coral is not elevated and reflects the influence of ambient water with a r14C signature equivalent to the 

surrounding dissolved inorganic carbon pool. 

Figure 4. Hawaiian black coral Leiopathes annosa. This 

species includes the oldest known coral. Photo credit: Hawaii 

Undersea Research Laboratory (HURL). 

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303

anthropogenic bomb-radiocarbon signal from 

the testing of thermonuclear devices in the 

1950s and 1960s. This method is sometimes 

referred to as bomb radiocarbon dating 

(Figure 3; Roark et al. 2005, Sherwood et 

al. 2005), which also has applications in the 

validation of fish age (Andrews et al. 2012).

II.1. Black Coral

Black coral (Figure 4) represent a group 

of organisms that are some of the deepest 

dwelling and longest-lived species. Several 

species live in U.S. territorial waters and 

most have been long lived (centuries to 

millennia). Wagner et al. (2012) summarized 

information on black coral growth and 

longevity. Longevity is particularly important 

within the black coral fishery (Parrish et al. 

and Wagner et al., this volume) because age 

and growth rates are needed to determine 

sustainable yields. These typically shallower 

dwelling species appear to be faster growing 

than deeper species and growth rates have 

been measured based on axial extension rates, 

growth rings, radiocarbon (14C) and lead-210 

dating (Grigg 1976, Roark et al. 2006, Love et 

al. 2007, Risk et al. 2009). Estimated longevities 

for sampled Antipathes spp. ranged from ~ 12 

- 140 years (Wagner et al. 2012).

In contrast, deepwater black corals from 

Hawaii (Leiopathes annosa, reported as 

Leiopathes spp.) were found to have a potential 

lifespan in excess of 4000 years with a radial 

growth rate of less than 10 μm·yr-1 (Roark et 

al. 2006, 2009). Similar estimated growth rates 

and longevities (400 – 2100 years) have been 

reported for Leiopathes from the Gulf of Mexico 

(Prouty et al. 2011) and the southeastern 

United States (Williams et al. 2007), with 

variable growth rates documented from 

populations in the Azores (Carreiro-Silva et al. 

2013). 

II.2. Gold Coral

The proteinaceous colonial parazoanthids 

(formerly known as Gerardia spp.), commonly 

referred to as gold corals, have similar 

longevity to that of Leiopathes. Numerous 

specimens (n = 23) of the Hawaiian gold 

coral (Kulamanamana haumeaae) (Figure 5) 

dated by radiocarbon show lifespans up to 

2740 years with an average radial growth 

rate of 41 ± 20 μm·yr-1 (Roark et al. 2006, 

2009, Parish and Roark 2009). These results 

are not in agreement with shorter lifespan 

estimates (max age ~70 years) and faster radial 

growth rates (~1 mm·yr-1) from presumed 

annual growth rings (Grigg 1974). A gold 

coral (identified as Gerardia sp.) specimen 

from Little Bahamas Bank in the Atlantic 

Ocean dated by amino acid dating (~250 

±70 years; Goodfriend et al. 1997) and by 

radiocarbon (1800 ±300 years; Druffel et al. 

1995) also revealed similar age and growth 

discrepancies. Using radiocarbon and stable 

isotopes (d13C, d15N), Roark et al. (2009) 

showed that K. haumeaae are feeding almost 

exclusively on recently transported particulate 

organic carbon from surface waters and that 

skeletal growth utilizes this carbon. An in 

Figure 5. The Hawaiian gold coral 

Kulamanamana haumeaae. Photo credit: HURL.

AGE, GROWTH RATES, AND PALEOCLIMATE STUDIES IN DEEP-SEA CORALS OF THE U.S.

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304

situ tagging study by Parrish and Roark (2009) 

observed no measurable change in the size of 

Hawaiian gold coral colonies over nine years, 

which is consistent with the slow growth, 

long-lived age estimates using radiocarbon 

dating. Based on radiocarbon results, and 

counter indicative to sclerochronology 

estimates, the Western Pacific Fishery 

Management Council implemented a 5-year 

moratorium on gold coral harvesting in 2008, 

which was subsequently extended through 

2018.

II.3. Precious Coral

Corals in the Family Coralliidae (e.g., pink and 

red corals in the genus Corallium) have been a 

historical part of the precious coral (Figure 6)

fishery in many parts of the world, primarily 

in the Mediterranean and western north 

Pacific (Bruckner and Roberts 2009). Most of 

these fisheries have been strongly impacted 

by fishing effort that exceeds sustainable 

growth. Age estimates based on a variety of 

petrographic (carbonate structure) and organic 

band counting are generally consistent with 

ages estimated using radiometric techniques. 

Colony age estimates range between 50 and 

180 years, with radial growth rates less than 

1 mm·yr-1 (Druffel et al. 1990, Marschal et al. 

2004, Andrews et al. 2005, Roark et al. 2006, 

Luan et al. 2013). In addition, deeper water 

Corallium species appear to grow more slowly 

than shallower species (Roberts et al. 2009), 

making them more vulnerable to exploitation. 

This has led to cautionary measures in some 

regions where deep-water Corallium is known 

to exist (DeVogelaere et al. 2005).

II.4. Bamboo & Other Octocoral

Radiocarbon and bomb-radiocarbon dating 

of living bamboo corals (Family Isididae) 

(Figure 7) range from 50 to 420 years and 

radial growth rates ranging 50 to 100 μm·yr-1

(Roark et al. 2005, Sherwood et al. 2009, Hill 

et al. 2011, Sinclair et al. 2011, Thresher et al. 

2011, Farmer et al. 2015). Based on the timely 

response of the bomb radiocarbon signal, it is 

likely that there is a correspondence of growth 

with surface water productivity. A recent 

innovation in lead-210 dating using a higher 

Figure 6. The Hawaiian precious coral Corallium 

laauense (= Hemicorallium laauense). In the 

past, this species has been harvested for jewelry in 

Hawaii. Photo credit: HURL.

Figure 7. The bamboo coral, Isidella tentaculum, 

from Giacomini Seamount in the Gulf of Alaska. 

Photo credit: NOAA and the Woods Hole 

Oceanographic Institution Alvin Group.

AGE, GROWTH RATES, AND PALEOCLIMATE STUDIES IN DEEP-SEA CORALS OF THE U.S.

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305

resolution approach provided age estimates 

of 50 to 100 years for bamboo coral in the NE 

Pacific (Andrews et al. 2009). Given that radial 

growth rates for this group appear to be less 

than 200 μm·yr-1, this group is susceptible to 

disturbance and removal because recovery 

would take decades. 

Red tree corals (Family Primnoidae) can reach 

over 2 m in height and are among the most 

important habitat-forming gorgonian corals in 

the Northeast Pacific and Northwest Atlantic. 

Maximum ages of live-collected red tree coral 

specimens from Eastern Canada (Primnoa 

resedaeformis) were ~ 78 -100 years (Sherwood 

et al. 2005, Sherwood and Edinger 2009) and 

from the Northeast Pacific (Primnoa pacifica) 

were ~ 119-185 years (Andrews et al. 2002, 

Aranha et al. 2014). Estimated average radial 

growth rates of P. pacifica of 320-360 μm·yr-1

were faster than those of the sister species, 

P. resedaeformis, in the Atlantic (Aranha et

al. 2014). Sub-fossil (i.e., dead corals whose

remains are not fully fossilized) specimens of

P. resedaeformis were estimated to be at least

700 years old (Sherwood et al. 2006).

Ages and radial growth rates of other 

deepwater gorgonian corals have been 

reported for Muricella sp. (Family: 

Acanthogorgiidae; ~50 and 100 years; 100-200 

μm·yr-1) from the 100 m depth range in the 

tropical Pacific (Williams and Grottoli 2010); 

and from the Northeast Atlantic, Paramuricea

sp. (Family: Plexauridae; ~ 70 – 100 years; 92-

205 μm·yr-1; depth 814-850 m) and Paragorgia 

arborea (Family: Paragorgiidae; 80 years; 830 

μm·yr-1; depth 814-850 m) (Sherwood and 

Edinger 2009). From the Gulf of Mexico, 

Prouty et al. (2014b) reported life spans of over 

600 years for Paramuricea biscaya, with radial 

growth rates between 0.34 μm yr-1 and 14.20 

μm yr-1 and linear growth rates from 0.019 cm 

yr-1 to over 1 cm yr-1.

II.5. Stony Coral

A few species of scleractinian corals, especially 

Lophelia pertusa (Figure 8), Solenosmilia 

variabilis, and Oculina varicosa, (and to a lesser 

extent Enallopsammia rostrata, E. profunda, 

Madrepora oculata and Goniocorella dumosa) 

create deepwater coral reefs or bioherms. 

These massive and ancient structures can 

be up to 30 m high and 10’s of kilometers 

in length (Reed 2004, Roberts et al. 2009). 

Lophelia reefs from lower latitudes, like those 

off of NW-Africa, the Mid-Atlantic Ridge 

and the Western Mediterranean Sea, indicate 

their structural growth represents 50,000 

years of accumulation, according to U/Th 

dating (Schroder-Ritzrau et al. 2005). Higher 

latitude Lophelia carbonate mounds, appear 

to have gone through alternating cycles of 

accumulation and die off over periods of 

centuries that appear to be tied to changes 

in oceanographic conditions associated with 

glacial-interglacial cycles (Roberts et al. 

2009) and the North Atlantic sub-polar gyre 

(Douarin et al. 2013). Individual colonies of 

L. pertusa from bioherms in the northeastern

Figure 8. Stony coral Lophelia pertusa in the 

Gulf of Mexico. Photo credit: NOAA and Bureau 

of Ocean Energy Management Lophelia II Project.

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306

Atlantic exhibited axial growth rates of 5-26 

mm·yr-1 based on in situ measurements, 

aquaria observations, isotopic analyses and 

estimates of age inferred from growth on 

artificial structures (Mortensen and Rapp 

1998, Mortensen 2001, Gass and Roberts 2006). 

Results with transplanted L. pertusa in the 

Gulf of Mexico exhibited axial growth rates of 

up to 16 mm·yr-1 with new polyps exhibiting 

higher growth rates and more mature polyps 

at rates of ≤5 mm·yr-1 (Brooke and Young 

2009). Observations of L. pertusa colonies on 

oil rigs and shipwrecks in the northern Gulf of 

Mexico (320-995 m depths) yielded minimum 

calculated growth rates ranging from 3.2 to 

32.3 mm·yr-1 (Larcom et al. 2014). 

Azooxanthellate Oculina varicosa has a 

geographically restricted distribution and 

forms reef-like structures in relatively shallow 

water (70 – 100 m). An axial branch growth 

rate of 16.1 mm·yr-1 was measured in-situ for 

O. varicosa at 80 m depth (Reed 1981). Similar

axial growth rates were found for Madrepora

oculata (14.4 ± 1.1 mm·yr-1) using lead-210

dating for samples collected off Norway

(Sabatier et al. 2011).

Solenosmilia variabilis is the dominant reefbuilding coral on seamounts in the southwest 

Pacific, where it occurs at depths significantly 

deeper than the Northern hemisphere L. 

pertusa reefs. Fallon et al. (2014) reported 

growth rates of from 0.84 – 1.25 mm·yr-1 linear 

extension for colonies collected between 958 

and 1,454 m. The authors estimated a coral 

accumulation rate of ~ 0.27 mm·yr-1, indicating 

that recovery from trawl damage would likely 

be extremely slow. Neil et al. (2011) found 

similar linear growth rates for S. variabilis in 

on Chatham Rise in New Zealand (ranging 

from 0.3 to 1.3 mm·yr-1) and estimated that it 

could take 380 to 1,700 years for colonies to 

grow to a maximum height of 1 m. 

Enallopsammia rostrata (Family

Dendrophylliidae) is an aborescent stony coral

that creates massive dendritic colonies up to

1 m wide and 0.5 m tall. Uranium-thorium

dating of specimens from the Line Islands

(Houlbreque et al. 2010), and lead-210 dating

of a single North Atlantic specimen, have

documented longevity up to ~600 years with

axial extension rates of 5 mm·yr-1 a radial

growth rate of ~0.07 mm·yr-1 near its base

(Adkins et al. 2004). The structure of E. rostrata

is prone to ‘shedding’ (natural limb loss) and

as a consequence accurate estimates of axial

growth rates are difficult to determine.

Other members of the stony corals have

different growth structures. The solitary cup

coral Desmophyllum dianthus has a slow axial

extension rate of 0.5-2.0 mm·yr-1 based on lead210, radiocarbon, and U/Th dating techniques

(Cheng et al. 2000, Adkins et al. 2002, 2004).

III. Paleoclimate and

Paleoenvironmental Studies

using Deep-Sea Corals

The utility of deep-sea corals in understanding 

past climate variability is closely tied to 

accurately dating the corals specimens 

(Robinson et al. 2014). Equally important 

is the development of environmental 

proxies for the temperature and elemental 

composition of the water in which the coral 

grew. As such, a fundamental goal in deepsea coral paleoclimate research has been to 

determine how environmental conditions 

are captured in the coral skeleton records. 

Both solitary and colonial, as well as calcitic 

and proteinaceous deep-sea coral species, 

are being used for climate change studies. 

Increased geographic distribution of sampling, 

coupled with advances in micro-analytical 

sampling techniques and recognition of novel 

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biogeochemical proxies, have advanced the 

field and are providing insights on climate 

variability at century to millennial time-scales.

III.1. Paleothermometry

Deep ocean circulation plays a vital role 

in modulating and stabilizing the Earth’s 

climate system, because the deep ocean 

stores and transports heat. Using radiocarbon 

captured within the skeletons of deep-sea 

corals, researchers have examined how the 

ocean has behaved in the past, in terms of 

rates of circulation and carbon exchange 

between different water bodies (e.g., Mangini 

et al. 1998, Frank et al. 2004). Evidence from 

deep-sea coral records indicate that the deep 

ocean circulation can change abruptly, on 

timescales as short as 10 years (Adkins et al. 

1998, Eltgroth et al. 2006). Such changes may 

have a major impact on global temperatures 

(Robinson et al. 2005), atmospheric carbon 

concentrations (Burke and Robinson 2012), 

and deep-sea organisms (Sutherland et 

al. 2012). Recent studies also indicate that 

tracers, such as the isotopic composition of 

neodymium from living and fossil deep-sea 

coral species (L. pertusa, D. dianthus and M. 

oculata), may have value in reconstructing 

ocean circulation patterns, particularly those 

operating during the last 10,000 years (van 

de Flierdt et al. 2006, 2010, Copard et al. 2012, 

Lopez Correa et al. 2012, Montero-Serrano et 

al. 2013).

To understand historical climate and 

ocean circulation changes, it is critical to 

reconstruct changes in seawater temperature. 

Ocean temperature reconstructions are 

challenging because biological processes 

also affect skeletal chemical compositions. 

The chemical composition of the skeleton is 

thus controlled by two main factors: 1) the 

external environment (e.g., temperature), 

and 2) biological activity (referred to as ‘vital’

effects). In deep-sea corals, these vital effects

may be larger than environmental controls

in elemental tracers, such as oxygen isotopes

(d18O) or trace metal ratios (e.g., Mg/Ca), which

are typically used as temperature proxies in

other marine organisms. However, by taking

an average of multiple analyses it is possible

to calculate environmental temperatures from

single specimens based on direct and indirect

comparisons (Smith et al. 2000, Adkins et al.

2003, Hill et al. 2011). Additional methods that

show promise for temperature reconstruction

include the ratio of Mg/Li in coral skeletons

(Case et al. 2010, Montagna 2014) and

“clumped” carbon and oxygen isotopes that

act independently of vital effects (Thiagarajan

et al. 2011). One important caveat for all of

these methods is that calculated uncertainties

for the paleothermometry estimates can range

from 0.5 to 2°C. The utility of reconstructing

small-scale temperature changes in the deepsea is limited by the precision of the technique

as well as temporal precision and accuracy.

Efforts are underway by a number of research

groups to increase the precision and resolution

with which deep-water temperatures can be

reconstructed.

III.2. Nutrient and Trophic Level

Proxies

Proteinaceous deep-sea corals, such as black 

corals, gorgonians (e.g., Primnoa spp.), and 

the colonial zoanthids gold corals (formerly 

known as Gerardia sp.), derive their skeletal 

protein from recently exported particulate 

organic matter from the surface. From a 

compositional perspective, these corals are 

somewhat analogous to sediment traps, 

integrating the geochemical signature of 

recently exported organic matter into their 

slow growing skeletal structure. Useful 

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geochemical signatures captured in skeletal 

protein include radiocarbon and stable carbon 

(δ13C) and nitrogen (δ15N) isotopes, which can 

be used to reconstruct nutrient sources and 

cycling as well as food web (trophic) dynamics 

through time. Elemental composition, such as 

phosphorus and barium, in both stony corals 

and gorgonians, can also be used in studies of 

seawater nutrients variability.

III.2.i – Nitrogen

With sufficient understanding of the nitrogen 

dynamics of a particular region, including 

the δ15N of sinking and suspended particulate 

organic matter, skeletal δ15N may be used 

in reconstructions of local trophic and/or 

nutrient dynamics. In the Gulf of Mexico and 

the South Atlantic Bight, marked increases 

over the past 75 years in coral skeletal δ15N 

exceed 3‰, indicating there has been a higher 

contribution of terrestrial effluent to the deep 

sea (Williams et al. 2007). In the western 

tropical Pacific, multi-decadal decreases in 

δ15N values from specimens at the base of the 

euphotic zone suggest a gradually shallowing 

of the nutricline (nutrient gradient) (Williams 

and Grotolli 2010). Off Tasmania, records of 

bulk δ15N indicate relatively stable nutrient 

and trophic conditions over the past 250 

years (Sherwood et al. 2009). In instances 

where the cause of skeletal stable isotopic 

variability is ambiguous, analysis of the δ15N 

of amino acids (δ15N-AA) has emerged as a 

powerful tool to separate and independently 

track the effects of source nutrients, trophic 

transfers and microbial activity (McCarthy et 

al. 2007, Prouty et al. 2014a). Using Primnoa 

resedaeformis samples from Nova Scotia, 

Canada, Sherwood et al. (2011) pioneered 

the application of d15N-AA to deep-sea 

corals, demonstrating a nutrient regime 

shift in the western North Atlantic since the 

1970s. These studies underscore the broad 

potential for proteinaceous deep-sea corals in

paleoceanographic studies linking nutrient

and trophic variability to changes in global

climate.

III.2.ii – Phosphorous and Barium

In addition to using isotopic studies from 

proteinaceous corals as proxies for nutrient 

input to the deep sea, two elements that have 

also been used are phosphorus and barium. 

Phosphorus is a key nutrient in global primary 

productivity. It is used by organisms at the 

ocean surface and concentrates at depth. As 

a result, variations in seawater phosphorus 

reflect changes in surface ocean biological 

production and cycling (Montagna et al. 

2006). As a proxy for seawater phosphate, 

phosphorus to calcium (P/Ca) ratios have 

been studied in the deep-sea coral D. dianthus

(Montagna et al. 2006, Anagnostou et al. 

2011). These global studies have shown a 

relationship between seawater phosphate 

and coral skeletal P/Ca ratios (Montagna et 

al. 2006, Anagnostou et al. 2011), suggesting 

that coral P/Ca has the potential to reconstruct 

variations in biological productivity on 

annual to decadal time-scales. Seawater 

barium, which also displays nutrient-like 

behavior in seawater, such that coral Ba/Ca 

ratios have the potential to trace the history 

of intermediate and deep-water refractory 

(slowly decomposed) nutrients, such as silica 

(Anagnostou et al. 2011, LaVigne et al. 2011, 

Sinclair et al. 2011).

III.3. Ocean Acidification

Ocean acidification is predicted to have 

profound implications for marine ecosystems 

partly because carbonate ions are an essential 

part of coral calcification. Changes to 

carbonate ion chemistry may particularly 

affect deep-water corals because carbonate 

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levels are already low (Guinotte et al. 2006, 

Turley et al. 2007, Thresher et al. 2011). 

Instrument and modeling studies indicate 

that seawater carbonate chemistry (carbon 

dioxide, pH, total dissolved inorganic carbon 

and alkalinity) is changing due to uptake of 

anthropogenic carbon dioxide (CO2

) from 

the atmosphere. In addition, it is likely that 

there were large changes during major climate 

events of the past. One way to examine the 

response of the ocean and deep-sea corals to 

large perturbations to the carbon cycle is to 

use geochemical proxies that record pH within 

the coral skeletons. 

Boron isotopes (d11B) in biogenic carbonates 

have been established as a proxy for seawater 

pH due to preferential uptake of the borate 

ion relative to boric acid as a function of pH 

(Vengosh et al. 1991, Hemming and Hanson 

1992). Experimental relationships between 

pH and d11B have been determined in both 

surface and deep-water scleractinian corals, 

indicating a relationship between carbonate 

d11B and seawater pH (Reynaud et al. 2004, 

Blamart et al. 2007, Maier et al. 2009, Trotter 

et al. 2011, McCulloch et al. 2012a). However, 

d11B values higher than predicted for seawater 

pH have been observed in these studies 

(e.g., Vengosh et al. 1991, Blamart et al. 2007, 

Trotter et al. 2011). In an effort to reconcile 

these observations, scientists have found that 

aragonitic (CaCO3

) forming corals (e.g., stony 

corals) are able to regulate internal pH at the 

site of calcification (McCulloch et al. 2012a, 

2012b). This may explain how some deep-sea 

corals can calcify at levels below the aragonite 

saturation horizon, which suggests deep-sea 

corals may be influenced less by decreasing 

seawater pH than originally thought 

(McCulloch et al. 2012a, 2012b, Hennige et al. 

2015). Therefore, one must be cognizant of the 

fact that the research is evolving and presently 

there are conflicting results.

III.4. Challenges

The geochemical archives derived from deepsea corals have enormous potential to help 

scientists decipher and describe changes 

and variability of paleoenvironmental and 

paleoceanographic conditions through time. 

However, it is inherently difficult to determine 

the accuracy of predictive relationships in 

the coral skeleton relative to the seawater 

environment from field data. Analyses of 

deep-water taxa are constrained by the 

challenges of collecting and sampling the 

organisms and sparse environmental data 

for use in validating relationships in ambient 

deep-sea environmental conditions. Deep-sea 

coral research is still in its infancy in terms of 

understanding variability between locations, 

depth strata, as well as taxonomic orders. For 

example, Thresher et al. (2010) suggest that 

adaptation to local conditions and hence a role 

for physiology at higher taxonomic levels may 

occur in deep-sea corals. Therefore challenges 

still exist to better constrain the speciesdependent effect and to identify the causes of 

inter-species differences and intra-colony age 

and growth variability (Carreiro-Silva et al. 

2013). 

Studies employing advanced micro-analytical 

techniques have confirmed that micrometer 

scale heterogeneity influences the geochemical 

signal in deep-sea corals. Recent studies on 

stony corals show that the main differences 

in minor/trace element compositions, stable 

isotopes and organic compounds occur 

between calcification centers and fibrous 

aragonite (Gagnon et al. 2007, RollionBard et al. 2009, López Correa et al. 2010). 

Biomineralization processes exerting a strong 

biological control on the skeletal formation can 

overwhelm signals caused by environmental 

conditions (Adkins et al. 2003, Gagnon et al. 

2007). Systematic studies of the isotopic and 

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310

trace metal variation in various biocarbonates, 

combined with detailed studies of deep-sea 

coral skeletal structure are critically needed 

to decipher the role of “vital effects” on the 

application of these proxies to paleoclimate 

studies.

IV. New Directions

There are several new directions being 

pursued in an attempt to successfully 

use deep-sea corals as reliable archives of 

seawater chemistry and oceanographic 

proxies to climate change research. Given 

that many common tracers in the carbonate 

skeletons of deep-sea corals are strongly 

affected by physiological processes during 

biomineralization (i.e., “vital effects”), there 

is a need to increase our knowledge of the 

characteristics and mechanisms of vital 

effects. In other words, how can we tease 

apart the strong biological fractionation 

that is superimposed on the environmental 

signal? Focused studies, such as addressing 

the question of how the fluid reservoir 

from which calcification occurs is linked to 

the surrounding seawater, are becoming 

increasingly important as we refine 

our understanding of the various coral 

calcification models and strategies to reduce 

bias from vital effects. The introduction of 

micro-analytical sampling techniques, such as 

micromilling, ion microprobes, laser-ablation 

and highly focused synchrotron radiation, 

has made it possible to investigate coral 

intraskeletal variability of trace element and 

stable isotopic compositions directly related 

to the ultra-structure of the skeleton (RollionBard et al. 2009, Thresher et al. 2009, 2010, 

Case et al. 2010, López Correa et al. 2010, 

Sinclair et al. 2011). Therefore, we need to 

couple these high-resolution measurements to 

increase knowledge of present day processes 

affecting the coral communities.

As previously mentioned, from a 

compositional perspective, proteinaceous 

corals are somewhat analogous to sediment 

traps since they depend on surface-derived 

particulate organic matter. Therefore, age 

and growth studies should be accompanied 

by sediment trap studies that can clarify how 

the elemental and isotopic composition of 

particulate organic matter (i.e., food source) is 

captured or represented in the deep-sea coral 

skeletal chemistry. Additional focused studies 

should include the collection of appropriate 

coral specimens along with food sources and 

particulate and dissolved nutrients from the 

same location to clarify our understanding 

of feeding habits, prey type, timing and 

seasonality, as well as the impact of microbial 

activity of food sources on geochemical 

signatures encoded in corals. Examination of 

new specimens from different nutrient and 

oceanographic regimes, integration of isotopic 

and trace elemental geochemical signals, and 

use of new techniques such as the δ15N of 

skeletal amino acids will aid in understanding 

past nutrient and trophic dynamics in the 

oceans, including reconstructions of source 

nitrogen variability and phytoplankton 

trophic and community structure.

A paramount challenge to the application 

and utilization of paleoenvironmental 

reconstructions using deep-sea corals 

is the precision, fidelity, and resolution 

of independently derived age-models 

(e.g., Komugabe et al. 2014), regardless of 

whether they are radiometric or based on 

sclerochronology. A particularly challenging 

time period is the ‘near instrumental’ period 

of the last several hundred years where 

radiocarbon is insensitive and sample 

size requirements make Th/U, in general, 

untenable for a well resolved chronology. 

Anthropogenic tracers with known or 

reconstructed emission histories (e.g., Pb, 

Pb-isotopes, As, Cd, Hg) and proxies of 

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disturbance events of known age may provide 

regional tie points (e.g., Schuster et al. 2002, 

Kelly et al. 2009) that could be coupled to 

radiometric chronologies. For example, 

Andrews et al. (2009) applied a refined lead210 dating technique to yield relatively highprecision growth rate and age determinations 

for bamboo corals living during the last 100 

years. Inter-disciplinary studies that use 

multiple chronological approaches with crossvalidation of sclerochronological features are 

also necessary. Validation of the timing and 

cause of sclerochronological features have the 

potential to refine chronologies. Advances in 

deep-sea coral research continue to highlight 

the growing importance of deep-sea corals as 

reliable marine archives of climate change and 

other environmental cycles. Like their shallowdwelling counterparts, deep-sea corals have 

also proven to be important archives of past 

ocean variability. While challenges still exist 

to providing reliable, reproducible records of 

climate variability, recent advances provide new 

opportunities and directions to close this gap.

Information on growth rate and life span of 

deep-sea corals is essential for conservation 

and management because the life history of 

these prominent organisms is either poorly 

understood or unknown. Validated age 

and growth of these organisms is the most 

fundamental information on susceptibility 

to disturbance or removal. Early estimates 

of age and longevity have been roughly 

determined, but more advanced techniques 

hold promise in not only determining 

longevity, but variations in growth through 

ontogeny (Andrews et al. 2009, Roark et al. 

2009, Carriero-Silva et al. 2013). Assessment 

of the vulnerability of these long-lived, 

habitat-forming organisms to both natural and 

anthropogenic perturbations is of paramount 

importance because recovery may involve 

life history aspects that are not currently 

considered. Once some or many of these

coral species are lost to damage or removal,

many are not likely to return within our

lifetime. Even in terms of minor damage, some

arborescent corals have shown little to no

recovery in nearly a decade of no disturbance

(Krieger 2002, Williams et al. 2010). In

addition, there is evidence that energy may

be focused on regenerative growth within the

broken colony in lieu of reproductive effort

(Waller and Tyler 2005). For the longest-lived

members of the deep-sea corals, perhaps it is

prudent to take on a perspective of value in

terms of world heritage, analogous to ancient

terrestrial forests.

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• • •
                         

           

Euplectella aspergillum glass sponge in the northwestern Gulf of Mexico. Courtesy of the NOAA

Office of Ocean Exploration and Research.

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

TEXAS TO THE FLORIDA STRAITS

  

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

STATE OF DEEP-SEA 

CORAL AND SPONGE 

ECOSYSTEMS IN THE 

GULF OF MEXICO 

REGION: TEXAS TO 

THE FLORIDA STRAITS 

Gregory S. 

Boland1*, 

Peter J. Etnoyer2, 

Charles R. 

Fisher3, and 

Emma L. 

Hickerson4

  

1Division of

Environmental Sciences,

Bureau of Ocean Energy

Management, Sterling, VA

* Corresponding Author:

Gregory.Boland@boem.gov  

2 NOAA Center for Coastal

Monitoring and

Assessment, National

Centers for Coastal Ocean

Science, Charleston, SC  

3Department of Biology,

The Pennsylvania State

University, State College,

PA

4 NOAA Flower Garden

Banks National Marine

Sanctuary, Galveston, TX

I. Introduction

This chapter provides an update and summary of the current state

of knowledge of deep (> 50 m) azooxanthellate coral and sponge

communities on hard‐bottom habitats in the northern Gulf of

Mexico (GoM) from the U.S.‐Mexico border to the Florida Straits.

This is an update since the first State of Deep‐Sea Coral Ecosystems

of the U.S. report (Lumsden et al. 2007) on the GoM by Brooke and

Schroeder (2007)1. The chapter will focus on new material but also

touch on some additional background information important in the

context of deep corals and sponges not presented in 2007.

1The 2007 GoM report (Brooke and Schroeder 2007) also included information on the Florida Straits up to Biscayne Bay on the east coast of

Florida. This geographic area is covered in the Southeast U.S. chapter of the current volume.

The GoM is unique in many ways, partly due to unusual geological

history and resulting physiographic features that provide substrate

for deep corals and sponges. The deep Gulf, from the outer portions

of the continental shelf near 150 m depth across the continental

slope to some of the deepest portions near 3,000 m depth, is

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

TEXAS TO THE FLORIDA STRAITS

  

Figure 1. Multibeam bathymetry of the northern Gulf of Mexico with depth shading of continental slope below

100 m showing the complexity of the slope. Created from industry‐acquired 3‐D seismic data processed at 50

foot bin size by the Bureau of Ocean Energy Management. Inset shows the Sigsbee Escarpment.

generally considered one of the most

geologically complex continental slopes in the

world (Figure 1). The geological complexity of

the GoM supports a high diversity of deep

corals, each adapted to different environmental

conditions (described broadly in Roberts et al.

2009). The vast majority (> 99%) of the substrate

in the central and western portion of the deep

northern GoM is composed of soft sediments in

mixtures of fine sand, silt and clay (Rowe and

Kennicutt 2009). Sea pens, cup corals and

bamboo corals can occur in soft sediments,

occasionally in high abundance over a large

area (Gallaway et al. 1988, Rowe and Kennicutt

2009). However, the highest diversity of large

structure‐forming coral and sponge epifauna

tends to occur on hard bottom. In the

mesophotic zone (30‐150 m), some of the

limited hard substrate is of biogenic origin (e.g.

drowned Pleistocene coral reefs). Many other

areas on the continental shelf are influenced by

movement of underlying salt deposits that can

raise the seabed to form banks or mounds

where in some cases, such as the Flower

Garden Banks, diverse zooxanthellate coral

communities can develop. In one location, the

only known example, basalt spires form a

volcanic chimney that is exposed at Alderdice

Bank. Hard‐bottom habitats below 200‐300 m

are primarily the result of diapirism of Jurassic‐

age salt associated with trapping and migration

of hydrocarbons.

Salt diapirism or ‘salt tectonics’ produced

hundreds of banks and mounds along the

continental slope in the northern region (west

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

TEXAS TO THE FLORIDA STRAITS

  

of the Florida Escarpment), because salts

typically are buoyant relative to overlying

sediments. Fissures in sediment overlying salt

domes can allow trapped hydrocarbons to seep

upwards to the seafloor which become a food

source for bacterial chemosynthesis at the seep

interface resulting in deposition of authigenic

(generated in place) carbonates which can form

large rocky features with moderate relief and

patchy distribution. Carbonates either develop

above the soft bottom substrate or become

exposed over time. Carbonate structures

associated with former and extant hydrocarbon

seeps provide substantial habitat for deep

corals and sponges in the northern GoM shelf

and slope region (CSA 2007, Brooks et al. 2016).

In contrast, the Florida platform and

escarpment along the eastern Gulf were formed

by deposition and consolidation of sediments

created by a variety of mechanisms including

deposition by carbonate‐producing organisms

such as corals and mollusks (Hine 2009). The

Bureau of Ocean Energy Management (BOEM)

subdivides the GoM into three operational

areas; Western, Central and Eastern (termed

Planning Areas), each with different levels of

industrial development and agency mission‐

oriented research investment (although many

studies are ecosystem‐based and include Gulf‐

wide locations and objectives). The northern

central GoM (Central Planning Area) is

historically the most active for research and

exploration. Due to a moratorium on drilling

offshore, the eastern GoM is less explored.

Since the last State of Deep‐Sea Coral

Ecosystems report (Brooke and Schroeder 2007)

there has been an increase in exploration and

research of the deep coral and sponge

assemblages of the West Florida shelf and

slope. Although data are limited, explorations

to date suggest that the extent of hard‐bottom

habitat is large and the abundance and

diversity of corals and sponges is high.

II. New Exploration and

Research

Research on deep corals in the GoM has

intensified substantially over the last decade.

Since 2007, at least 52 research cruises have

taken place in this region (Tables 1‐2). A

number of large multidisciplinary interagency

projects have greatly expanded the number of

known deep coral habitats in the GoM, and

increased our knowledge of their distribution

and community structure, as well as dispersal,

growth and reproduction of key species.

Federally sponsored explorations of the deep

GoM were planned for 2008‐2012, but the 2010

GoM oil spill resulting from the blowout of the

Macondo well and the sinking of Deepwater

Horizon (DWH) drilling platform at a depth of

1,522 m was a major environmental event with

profound ramifications for research and

management. One effect of the spill was a

dramatic increase in research effort as part of

the federally mandated Natural Resource

Damage Assessment (NRDA) for DWH. NRDA

conducted multi‐year investigations of deep

corals in the mesophotic zone (depth range 65‐

90 m), and in the deep‐sea (depth 1500 m and

more). These studies included investigations of

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Date Vessel Underwater 

vehicle Sites Notes

Mar. 2007 

Mesophotic Coral Habitat Exploration by Flower Garden Banks National Marine Sanctuary 

Horseshoe, Rankin/Bright, EFG and M/V Carolyn Choest HOV Argus/NR1 Secrets of the Gulf Expedition - Dr. Robert Ballard WFG, 

May 2009 R/V Manta ROV Phantom Stetson, McGrail and MacNeil Banks 

May 2010 R/V Manta ROV Phantom EFG and WFG NCCOS 

May 2011 R/V Manta ROV Phantom EFG and WFG NCCOS 

Aug.-Sep. 2012 R/V Manta ROV Phantom EFG and WFG NCCOS 

Oct. 2013 R/V Manta ROV Mohawk EFG 

Jul. 2015 R/V Manta ROV Mohawk McGrail, Bright and Geyer Banks FAU/CIOERT/Voss Lab 

Jul. 2015 R/V Manta ROV Mohawk Stetson Bank Stetson Bank LTM 

Jul. 2015 

EFG, Horseshoe, Rankin, Bright, Rezak, 

R/V Manta ROV Mohawk Elvers, Sidner, Bouma, Bryant, Alderdice 

and Parker Banks 

Jun.-Jul. 2007 R/V Ronald H. Brown 

BOEM/NOAA OER; Chemo III 3rd Field Sampling Expedition 

Chemo III primary sites, GB697, AT320, ROV Jason II GC852, GB647, GC645, AC601 Spectacular Madrepora colonies at 1,400 m at 

GC852 

BOEM/NOAA OER; Lophelia II 

Sep.-Oct. 2008 R/V Nancy Foster ROV Sea Vision 4 shipwrecks, 13 coral sites 

Jun.-Jul. 2009 R/V Brooks McCall AUV Sentry 8 coral sites 

Aug.-Sep. 2009 

Oct.-Nov. 2010 

R/V Ronald H. Brown 

R/V Ronald H. Brown 

ROV Jason II 5 shipwrecks, 12 coral sites 

ROV Jason II 2 shipwrecks, 15 coral sites 

10 new coral sites 

5 new coral sites, first observation of corals 

impacted by Deepwater Horizon spill at block 

MC294. 

Jul. 2012 R/V Brooks McCall ROV Kraken II 5 deep-water oil and gas structures 4 surface structures, 1 subsea installation 

USGS Collaboration Expeditions for Lophelia II 

Oct. 2008 R/V Nancy Foster ROV Sea Vision VK826, 862/906, West Florida Shelf 

Sep. 2009 R/V Seward Johnson HOV Johnson Sea Link VK826, 862/906, West Florida Shelf 

Sep.-Oct. 2010 R/V Cape Hatteras ROV Kraken II VK826, 862/906, West Florida Shelf 

Oct. 2010 R/V Arctic Sunrise HOV Dual DeepWorker VK862/906 Lander deployment 

Oct.-Nov. 2011 R/V Ronald H. Brown ROV Jason II West Florida Shelf 

Oct.-Nov. 2011 R/V Tommy Munroe ROV Global Explorer VK862/906 Lander recovery 

Florida Shelf Edge Exploration (FloSEE); NOAA OER, Cooperative Institute for Ocean Exploration, Research and Technology 

Jul. 2010 R/V Seward Johnson HOV Johnson Sea Link West Florida Shelf to Alabama 

Sep. 2011 R/V Nancy Foster ROV Kraken II West Florida Shelf to Alabama 

Ecosystem Impacts of Oil and Gas Inputs to the Gulf (ECOGIG); Ocean Exploration Trust 

Jul. 2013 E/V Nautilus ROV Hercules All Gulf

Jun. 2014 E/V Nautilus ROV Hercules All Gulf 

Apr.-May 2015 E/V Nautilus ROV Hercules All Gulf

BOEM study; Deep-Water Reconnaissance of Potentially Sensitive Biological Features 

Oct. 2011 R/V Manta ROV Phantom Horseshoe and 29 Fathom Banks 

Sep. 2012 R/V Manta ROV Phantom Rankin and 28 Fathom 

Sep. 2012 R/V Manta ROV Phantom 28 Fathom, Bright, Geyer 

Apr. 2013 R/V Manta ROV Phantom Alderdice and Rezak 

Apr.-May 2013 R/V Manta ROV Phantom Rezak, Bouma, Sonnier 

Jun. 2013 R/V Manta ROV Phantom Elvers and McGrail 

Jun. 2013 R/V Manta ROV Phantom Sidner and Parker 

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

Table 1. Research expeditions targeting deep‐sea coral and sponge habitats in the Gulf of Mexico since 2007.

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Date Vessel Underwater 

vehicle Sites Notes

Gulf of Mexico Exploration; NOAA OER 

Apr. 2012 E/V Okeanos Explorer ROV Deep Discoverer Northwest Gulf EX1202

May 2012 E/V Okeanos Explorer Florida Escarpment and Straits, Northeast ROV Deep Discoverer Gulf EX1203 

Apr. 2014 E/V Okeanos Explorer ROV Deep Discoverer All Gulf EX1402

Apr.-May 2014 R/V Atlantis II HOV Alvin 

Gulf of Mexico Exploration; Alvin

North central Gulf Science mission on deep coral and chemosynthetic 

community sites 

Schmidt Ocean Institute Exploration, Mapping and Long-Term Effects of the DWH Spill 

Aug. 2012 R/V Falkor ROV Global Explorer Northeast Gulf Deep-sea coral shakedown expedition 

Sep. 2012 R/V Falkor ROV Global Explorer Northwest Gulf, Texas Banks Mapping of South Texas Banks 

Nov. 2012 R/V Falkor ROV Global Explorer Northwest Gulf Long-term effects of DWH spill 

Mar. 2012 R/V Maria S. Merian 

General Exploration; Ocean Exploration Trust 

ROV Cherokee West and southwest Florida slope 

General cold-water coral ecosystem exploration 

including other areas in southern GoM and outside 

GoM 

Jul. 2014 E/V Nautilus ROV Hercules All Gulf Exploring the unknown Gulf of Mexico 

                       

Date Vessel Underwater 

Vehicle Sites Notes 

Jul. 2010 

Aug. 2010 

Nov. 2010 

Mar. 2011 

Apr. 2011 

Aug. 2011 

Oct. 2011 

Jun.-Jul. 2014 

R/V Nancy Foster 

R/V Nancy Foster 

R/V Brooks McCall 

HOS Sweetwater 

R/V McArthur 

M/V Holiday Choest 

M/V Holiday Choest 

R/V Walton Smith 

Natural Resources Damage Assessment Cruises 

ROV Global Explorer VK sites and MC751 

ROV Global Explorer Alabama Alps, Roughtongue Reef, Coral Trees Reef 

Drift Camera Well site and MC334

Industry ROV MC294 and MC118 

AUV Sentry Well site vicinity 

Alabama Alps, Roughtongue Reef, Yellowtail Reef, UHD Shilling Robotics Coral Trees Reef, Madison-Swanson South Reef 

UHD Shilling Robotics previously discovered sites 

Alabama Alps, Roughtongue Reef, Yellowtail Reef, 

ROV Global Explorer Coral Trees Reef, Madison-Swanson South Reef, 

Madison-Swanson North Reef 

Mesophotic reef assessment 

MC344 corals discovered

 

5 new sites found 

Mesophotic reef assessment 

Mesophotic reef assessment 

ECOGIG Expeditions (Ecosystem Impacts of Oil and Gas Inputs to the Gulf - Part of Gulf of Mexico Research Initiative) 

Nov. 2012 R/V Falkor ROV GE Impacted and control sites Additional exploration 

Jun. 2013 R/V Nautilus ROV Hercules Impacted and control sites Additional exploration 

Jun.-Jul. 2014 R/V Nautilus ROV Hercules Impacted and control sites Additional exploration 

Dec. 2010 

Mar. 2012 

R/V Atlantis 

E/V Okeanos Explorer 

Other Cruises Associated with Damage Assessment 

HOV Alvin, AUV Sentry Return to MC294 and MC344 

ROV Deep Discoverer Portion of virtual cruise, previous coral sites Additional exploration 

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

Table 1. continued.

Table 2. Research expeditions after Deepwater Horizon spill to assess deep‐water coral damages.

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

coral associated communities of fish, red crabs,

infaunal and epifaunal invertebrates in

surrounding soft sediments. Each of the

relevant major developments will be discussed

in more detail below. New deep coral research

since 2007 can be divided into three broad

categories.

• Continental slope studies; applied science

and exploration missions (Table 1);

• Continental shelf studies of mesophotic reef

habitats (Table 1);

• DWH‐related response and NRDA cruises

(Table 2).

II.1. Continental Slope Studies

II.1.i – Large federal interdisciplinary

studies 

Two large federal multidisciplinary deep‐water

studies have been completed since 2007,

sponsored by the National Oceanographic

Partnership Program (NOPP) and funded by

BOEM and the National Oceanic and

Atmospheric Administration (NOAA) through

the Office of Exploration and Research (OER).

The intention of these studies was to conduct

research that addresses the needs of BOEM,

which is charged with regulating the

development of oil and gas resources in the

deep GoM in an environmentally sound

manner. The exploratory mission and other

agency objectives of NOAA OER were also met.

These studies had complimentary projects

separately funded by United States Geological

Survey (USGS), and are briefly described

below:

 

a. Investigations of chemosynthetic

communities on the lower continental

slope of the GoM, (Chemo III): 2005-2009

This multidisciplinary study was initiated by

partners BOEM and NOAA OER in 2005 to

investigate both chemosynthetic and coral

communities at depths below 1,000 m. This

depth zone had received little research effort

because regional submergence facilities (e.g.,

the Johnson Sea Link submersibles) were limited

to ~1,000 m depth. Three major cruises were

conducted during the Chemo III study

including the use of the Alvin manned

submersible and the remotely operated vehicle

(ROV) Jason II. At most of the 13 primary study

sites there were observations of at least a few

large colonial cnidarians; however, only one

site (in lease block GC852) visited during this

study was found to have a diverse coral

community. This site included three species of

scleractinian structure‐forming corals

(Madrepora oculata, Enallopsammia rostrata and

Solenosmilia variabilis) at a depth of 1,435 m

(Brooks et al. 2014, Brooks et al. 2016).

In a related study, USGS partners conducted a

series of cruises (2006–2009) visiting a subset of

Chemo III stations. This research incorporated a

variety of additional research topics, including

tropho‐dynamics of fish populations and mid‐

water trawl collections above deep coral

communities to investigate connectivity. USGS

researchers also returned to a shallower deep

coral site within Viosca Knoll lease block 826

(VK826, 500‐700 m). This is a well‐known deep

coral site that was a primary focus during the

earlier Lophelia I research program (funded by

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b. Exploration and research of northern

GoM deep-water natural and artificial

hard-bottom habitats with emphasis on

coral communities: reefs, rigs and wrecks

(Lophelia II): 2008-2012

                 

           

             

               

           

         

             

           

               

           

             

                 

         

           

           

           

               

                   

             

       

             

         

        

                                                     

  

             

         

       

             

               

                

               

         

               

     

               

         

       

           

           

           

         

           

             

                 

         

               

                 

                 

               

       

           

         

       

     

        

             

             

           

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

BOEM) from 2004‐2007 (CSA 2007). The USGS

cruises included additional study sites at Viosca

Knoll, Atwater Canyon, Alaminos Canyon, and

Green Canyon (Demopoulos et al. 2010, Ross et.

al. 2012).

Lophelia II was the second of the BOEM and

NOAA OER studies on deep coral ecosystems

in the GoM, and this represented a continuation

and expansion of Lophelia I. As with the

previous BOEM studies, Lophelia II brought

together numerous collaborating groups and

again partnered with USGS to continue the

exploration and research of deep coral

communities in the deep GoM (Brooks et. al.

2016). Lophelia I spanning 2004‐2007 was

referenced in Brooke and Schroeder (2007) but

the report was final in July 2007 (CSA 2007).

The partnership between federal and academic

scientists included exploration and research at

three different types of deep‐water habitats:

natural reefs, shipwrecks and offshore energy

platforms in water depths ranging from 300 to

2,700 m. One primary goal of the study was to

develop a robust capability for predicting the

occurrence of rich cnidarian hard‐bottom

communities in the deep GoM, with emphasis

on scleractinian coral assemblages. In addition,

long‐term monitoring stations were established

2http://fl.biology.usgs.gov/DISCOVRE/gom.html

to study change over time along with

laboratory experiments to provide a more

comprehensive understanding of the processes

that control the occurrence and distribution of

Lophelia pertusa in the GoM (Larcom et al. 2014,

Lunden et al. 2014, Brooks et al. 2016).

A total of five Lophelia II cruises (eight

including USGS cruises) were completed

between 2008 and 2012 (Table 1). During these

cruises, researchers discovered new coral‐

dominated sites on the northern GoM slope and

the west Florida Escarpment, documented coral

colonization of shipwrecks and deep‐water

offshore platforms, and documented changes at

sites discovered during Lophelia I. The

shipwreck and platform components of the

Lophelia II study represented important

additions to the understanding of coral

habitats. Eight wreck sites were surveyed to

depths of over 2,100 m, in addition to those

described previously (Brooke and Schroeder

2007, Church et al. 2007). Many had extensive

coral cover (Larcom et al. 2014, Brooks et al.

2016). A total of 24 natural habitat study sites

were surveyed with an ROV during the project.

Community structure, biodiversity and

biogeography of corals and associates were

addressed by different techniques including

photographic transects, photo mosaics,

quantitative collections of community

components and molecular methods.

Lophelia II had strong collaboration with USGS

scientists, which was highlighted on the USGS

DISCOVRE Program website.

2 A series of

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

reports were produced summarizing the

Lophelia II findings beginning with a final

BOEM interim report in 2012 (Brooks et al.

2012) followed by the BOEM final report in

2016 (Brooks et al. 2016). Numerous blogs,

mission essays, images, video clips, and other

information were also made available from

expedition webpages for each cruise in 2008,

2009, 2010, and 2012 on the NOAA Ocean

Explorer website. The USGS final report is in

revision (Demopoulos et al. in revision) but

other publications were completed and are

available (e.g., Morrison et al. 2011, Prouty et al.

2016).

II.1.ii – Other deep-sea expeditions in the

GoM since 2007

In addition to the Lophelia II studies, several

other deep coral explorations have taken place

in the Gulf since 2007 (Table 1). The Johnson‐

Sea‐Link II manned submersible supported the

Florida Shelf Edge Expedition (FloSEE) in 2010

and followed up one year later aboard NOAA

ship Nancy Foster in 2011 deploying the Kraken

II ROV. The R/V Seward Johnson and Johnson‐

Sea‐Link II submersible concluded its legacy in

the GoM in 2010. Harbor Branch

Oceanographic Institute (HBOI) operated the

vessel since 1985, but the vessel was sold in

2010. The NOAA ship Okeanos Explorer (2012

and 2014) and Ocean Exploration Trust’s E/V

Nautilus (2013, 2014, and 2015) engaged in

telepresence cruises that provided shore‐based

observers real‐time remote access to images of

deep coral habitats using fiber‐optic ROV feeds

from the GoM seafloor.

Two new research platforms had maiden dives

to deep coral and sponge habitats in the GoM.

The R/V Falkor (Schmidt Ocean Institute)

conducted ROV dives during its Deep‐Sea

Coral Shakedown Expedition in August 2012,

with subsequent cruises to the South Texas

Banks (September 2012) and other previously

explored sites (November 2012). The R/V Falkor

and E/V Nautilus both supported deep coral

investigations by the Ecosystem Impacts of Oil

and Gas Inputs to the Gulf (ECOGIG)

consortium funded through the Gulf of Mexico

Research Initiative. The Alvin II submersible

(Woods Hole Oceanographic Institution) also

conducted its first science dives in the GoM

(March – June 2014, dives no. 4679‐4737) on a

science verification cruise focusing on deep‐

water seep habitats in the northwestern GoM

and the Florida Escarpment, funded by the

National Science Foundation (NSF).

II.1.iii – Highlights of research findings from

the continental slope

Valuable new information has become available

from the many field and experimental studies,

including data on the survival and growth rate

of L. pertusa (Brooke and Young 2009, Larcom et

al. 2014, Lunden et al. 2014), the extreme

longevity of black corals in the genus Leiopathes

(Prouty et al. 2011) and gorgonian octocorals in

the genus Paramuricea (Prouty et al. 2016), as

well as the vulnerability of these corals to

pollution and disturbance (Hsing et al. 2013,

DeLeo et al. 2016). The branching stony coral L.

pertusa was found to grow in thickets on

offshore energy platforms and shipwrecks with

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

linear extension rates of 1‐2 cm/yr on average

(Larcom et al. 2014). Large Paramuricea and

Leiopathes colonies (> 1 m) between 300‐1,500 m

depth were found to reach ages of 660 years for

Paramuricea and as old as 2,100 years for

Leiopathes (Prouty et al. 2016). The growth rates

of these corals may depend on their proximity

to major nutrient resources like the Mississippi

River (Prouty et al. 2011) or the Loop Current.

Because these corals are heterotrophic

suspension feeders, it is worth noting that

resource availability is likely to vary on finer

scales as well, for example on different sides of

a mound or canyon feature which may also

contribute to different growth rates.

A few long‐term monitoring studies measured

bottom currents over extended (one year) and

short‐term periods in the northern Gulf. Long‐

term studies showed maximum bottom

currents peaked at only 22.6 cm/s at the 1,420 m

GC852 site, but a substantial peak of 60 cm/s

occurred at the shallower 440 m site at Viosca

Knoll (VK826, Brooks et al. 2016). Average

current speed at VK826 was 8 cm/s, but there

were periodic shifts in the current from west to

east, corresponding with a decrease in

temperature and salinity (Mienis et al. 2012)

that suggest two alternating water masses

impinging on the reef. Sediment traps were

deployed at VK826 as well to identify and

quantify sources of nutrition to the suspension

feeding benthic assemblage. Two food

pathways to Lophelia were identified from trap

samples: sinking organic matter (phytodetritus)

from the Mississippi River and vertically

migrating zooplankton (Mienis et al. 2012).

Significant contributions were made regarding

the associations and limits of temperature,

salinity, dissolved oxygen and aragonite

saturation states for L. pertusa at natural reefs

(Davies et al. 2010, Davies and Guinotte 2011,

Mienis et al. 2012, Lunden et al. 2013) and in

aquaria (Brooks et al. 2012, Lunden et al. 2014,

Brooks et al. 2016). The environmental

conditions at natural reefs were similar to those

recorded for Lophelia reefs in the eastern North

Atlantic, though dissolved oxygen and density

were lower in the GoM (Davies et al. 2010).

Interestingly, the aragonite saturation state was

found to be lower at sites where L. pertusa was

present than at sites where it was absent,

suggesting that this factor was likely not an

exclusive control of L. pertusa presence in the

deep GoM (Lunden et al. 2013). The thermal

tolerance of L. pertusa was also investigated.

Experimental studies suggest that a

temperature threshold occurs near 12 °C

(Brooks et al. 2016). These results will help

scientists and managers to understand the

effects of climate change on deep coral habitats

now and in the future.

II.2. Continental Shelf Studies of

Mesophotic Habitats

The mesophotic zone is a transition zone

between shallow reefs in the photic zone and

deep coral and sponge ecosystems in the

aphotic zone. These habitats occur between 30

and about 130 m depths on the continental shelf

(this report includes heterotrophic corals in this

zone at depths ≥ 50 m under the broad term

deep corals). Reefs occur on substrate related to

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

salt domes fossil structures and other hard

bottom features in the northern and eastern

GoM on the Florida Shelf. The total area of

mesophotic seafloor habitat in the GoM is vast,

estimated by Locker et al. (2010) to be 178,885

km2 (between 30‐100 m), an order of magnitude

larger than that depth zone of the U.S.

Caribbean or the main Hawaiian Islands, at

3,892 km2 and 3,299 km2, respectively. The GoM

continental shelf is broad and slopes gently,

allowing mesophotic ecosystems to occur

further offshore than in other parts of U.S.

waters. These reefs may be more isolated from

nearshore effects, like fishing intensity and

nutrient runoff (Locker et al. 2010).

Epicenters of new research in the GoM

mesophotic zone since 2007 were in three

primary regions: (1) the Northwestern GoM,

including the Flower Garden Banks National

Marine Sanctuary (FGBNMS) and the South

Texas Banks off the coast of Texas, (2) the

Pinnacle Trend in the northern Gulf off

Louisiana, Alabama and Mississippi, and (3)

Pulley Ridge in the southeastern Gulf off

Florida. Research on these ecosystems since the

late 1970s has focused on exploration of hard

bottom banks and mounds elevated above

surrounding soft‐bottom substrate in order to

understand the character and abundance of the

associated biological communities. Manned

submersibles (Bright and Rezak 1978, Reed et

al. 2005, 2006) and ROVs (Gittings et al. 1992,

Weaver et al. 2002, Schmahl and Hickerson

2006) were used to explore and characterize

dozens of pinnacles, banks and mounds in the

Gulf’s mesophotic zone, and most were

reported as good habitat for scleractinian,

antipatharian, stylasterid and gorgonian corals.

The purchase of a Forum Energy Technologies

Mohawk ROV by the National Marine

Sanctuary Foundation (NMSF) in 2013 has

increased the exploration and characterization

capabilities of the region. This science class

ROV is operated by the University of North

Carolina at Wilmington, Undersea Vehicle

Program through an NMSF agreement. The

vehicle is depth rated to 600 m and currently

outfitted for operations down to 365 m.

II.2.i – New research in the northwestern

GoM

The FGBNMS has conducted biological surveys

since 2001 on reefs and banks in the

Northwestern GoM to document the occurrence

of deep corals and sponges within and outside

the Sanctuary. The FGBNMS and partners have

conducted 17 ROV cruises to visit a total of 16

banks between 2007‐2015. Octocoral and black

corals are abundant and diverse components of

the benthic assemblages explored by these ROV

expeditions. Survey photos and videos since

2003 were analyzed as part of a doctoral

dissertation on octocorals (Etnoyer 2009) and a

master’s thesis on mesophotic black corals

(Nuttall 2013) at Texas A&M University. The

deep coral assemblages of the nearby South

Texas Banks were mapped, surveyed and

explored as well (Rodriguez 2015).

Additionally, Dr. Suzanne Fredericq from the

University of Louisiana – Lafayette, is

investigating algae populations on the

Northwestern GoM reefs and banks, including

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

comparisons of areas affected by DWH (Felder

et al. 2014). The information on the complexity

of shelf‐edge habitats from these surveys, and

the imagery data obtained by FGBNMS and

USGS, has helped to support management

planning and development of proposed

extensive numbers of features with significant

relief revealed by multibeam data that lie

outside No Activity Zones. This collaborative

project began in 2011 in partnership with

Louisiana Universities Marine Consortium,

FGBNMS, and the University of North Carolina

Figure 2. High‐resolution multibeam bathymetry map of the Bright and Rankin Banks showing complex

topography and features. Red shaded areas represent BOEM no activity zones above a depth of 85 m. (Images

courtesy of USGS, http://walrus.wr.usgs.gov/pacmaps/wg‐index.html).

sanctuary expansion plans. The investigations

described above and the multibeam surveys of

numerous GoM topographic features published

by Gardner et al. (1998, 2002; Figure 2)

prompted a broad new study awarded by

BOEM in 2011. The purpose was to investigate

the significance of features outside of existing

BOEM environmental protection areas called

No Activity Zones that prohibit all energy

development activities. Figure 2 illustrates

at Wilmington Undersea Vehicle Program. The

objective of this project was to survey outside

No Activity Zones and inside core‐biological

zones to investigate habitat and species

composition at these newly discovered

bathymetric features (Sammarco et al. 2016).

The first ROV cruises took place in October

2011, and continued through 2013 with two

cruises in September 2012, two cruises in April

2013 and two cruises in June 2013 (see Table 1).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

3http://www.ncddc.noaa.gov/website/google_maps/FGB/mapsFG

B.htm

Results of continued research will lead to more

refined adaptive management and protective

measures for these Potentially Sensitive

Biological Features including the potential

expansion of No Activity Zones.

In addition to the efforts by FGBNMS, NOAA’s

National Center for Coastal Ocean Science

(NCCOS) conducted quantitative surveys in the

mesophotic depth ranges of the Sanctuary. The

project was titled “Fish and Benthic

Communities of the FGBNMS: Science to

Support Sanctuary Management” (Clark et al.

2014). The Cooperative Institute for Ocean

Exploration, Research and Technology

(CIOERT) also initiated investigations of

hermatypic corals in the mesophotic depth

ranges at the FGBNMS and Bright and McGrail

Banks. In 2011, the NOAA ship Nancy Foster

mapped Elvers Bank over an area of 30 km2

.

Mapping has also taken place at Parker, Ewing,

and Claypile Banks by FGBNMS and Texas

A&M University. An online GIS‐based

mapping tool was developed to allow for virtual

exploration of these deep‐water habitats.

3

Catalogs of the major biological components are

available through this portal.

The FGBNMS is actively pursuing an expansion

plan that draws on these studies to support the

incorporation of additional mesophotic reefs

and banks in the northwest region. A draft

environmental impact statement addressing

sanctuary expansion was released on June 10,

2016, and presents a range of alternatives,

including a preferred alternative

recommending the addition of fifteen banks to

the Sanctuary, which would increase the

protected area from 56.21 miles2 to 383.19

miles2. This recommendation stemmed directly

from the mapping, exploration and

characterization activities in the region.

II.2.ii – New research in the Pinnacle Trend

region

In the Northeastern GoM, mesophotic reef

research has focused on the deep‐water reefs of

the Pinnacle Trend off Mississippi, Alabama

and Louisiana (Alabama Alps, Roughtongue

Reef and Yellowtail Reef) and on Coral Trees

Reef and Madison Swanson Marine Reserve

reefs south of the Florida panhandle. The

Pinnacle Trend reefs were below the oil slick

created by DWH spill for a period of several

weeks (Figure 3). These surveys commenced in

August 2010, three months after the spill,

following a period of little research activity on

deep corals and sponges in that area since 2003.

New studies reported on the density, health

and condition of gorgonian and black corals in

relation to days below the oil slick,

polyaromatic hydrocarbons (PAHs) in tissues

and sediments, as well as fishing gear impacts

(Silva et al. 2015, Etnoyer et al. 2016), in a 20‐

year retrospective analysis of ROV surveys.

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Figure 3. Map showing known deep‐sea coral aggregations (colored circles) in relation to the extent and

duration of the surface oil slick produced by the Deepwater Horizon blowout at Macondo wellhead in lease

MC252 (white triangle). Yellow circles represent mesophotic reefs at Alabama Alps Reef (AAR), Yellowtail

Reef (YTR) and Roughtongue Reef (RTR); green circles represent Lophelia pertusa reefs at Viosca Knoll; red

circles represent deep gorgonian sites near Macondo wellhead (MC) at 850‐1,850 m.

II.2.iii – New research in the GoM including

Pulley Ridge

Research since 2007 in the Eastern GoM has

focused on Pulley Ridge and Tortugas reefs,

and shelf‐edge habitats at Sticky Grounds,

Madison‐Swanson and Steamboat Lumps.

These are true mesophotic reefs of

zooxanthellate corals (Leptoseris and Agaricia) in

deep‐water to 90 m (Collier et al. 2015).

Harbor Branch Oceanographic Institute at

Florida Atlantic University (HBOI‐FAU) led the

Florida Shelf Edge Ecosystem (FloSEE) cruises

in 2010 and 2011 under the auspices of CIOERT

(Table 1). Results were detailed in reports by

Reed and Rogers (2011), Reed and Farrington

(2012), and Reed et al. (2012 a, b, c). One focus

of the FloSEE work was in response to DWH,

but the original intent of the effort was to

characterize poorly known mesophotic and

deep‐water habitats on the West Florida Shelf

using the Mohawk ROV, operated by UNCW‐

Undersea Vehicles Program.

Another project in the Eastern GoM mesophotic

zone was titled “Connectivity of the Pulley

Ridge – South Florida Coral Reef Ecosystem.”

Field expeditions took place in August 2012 and

2013 (Reed et al. 2014), sponsored by NOAA’s

NCCOS Center for Sponsored Coastal Ocean

Research (CSCOR) and NOAA OER. The 5‐

year, $5 million project focused on population

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connectivity, specifically the role that

mesophotic reefs may play in replenishing

corals, fish and other organisms in downstream

reefs of the Florida Keys and Dry Tortugas.

The studies were primarily concerned with

zooxanthellate corals and fishes in the

mesophotic zone, but deep‐water heterotrophic

octocorals were documented as well (Table 1).

II.3. Research associated with the

DWH Oil Spill

The DHW oil spill in April 2010 catalyzed a

large number of assessment and research

activities, including substantial new research on

deep corals. The total volume of oil entering the

GoM was approximately 651.85 million liters

(172,200,000 gallons) after subtracting the

volume of oil captured by various techniques

(McNutt et al. 2011). This represents the second

largest human‐caused oil spill into the

environment after the intentional 1991 Kuwait

oil field destruction. During the DWH spill, the

oil escaped at a depth of 1,500 m for a period of

100 days, and took the form of a submerged

plume and an oil slick that was present on the

sea surface and tracked by satellite for a period

of several weeks (Figure 3). Deep corals were

among the first benthic organisms targeted for

assessment and linked to the DWH spill.

Damaged corals were discovered in lease block

MC294, 11 km southwest of the Macondo well

site (White et al. 2012). The area was identified

as good potential for coral development by

BOEM and NRDA investigators using industry

3D seismic seabed anomaly data, and

discovered during the final dive of the

previously scheduled Lophelia II field sampling

cruise on November 2, 2010. The main area

reported in White et al. (2012) measured 10 x 12

m, and about 43 gorgonian octocorals,

primarily Paramuricea biscaya, were documented

in this area or within 10 m.

Impacts to gorgonian live tissue were observed

using high‐definition video from the Jason II

ROV, and a month later, from the manned

submersible Alvin. Injuries were observed to

86% of the corals. Analysis of hopanoid

petroleum biomarkers isolated from the brown

floc on the damaged corals provided strong

evidence that this material contained oil from

the Macondo well (White et al. 2012). A

calculation based on the age of the corals and

the odds of the impact happening coincidently

at this location and time (and not at any of the

other 20 coral sites visited during that time

period), yielded a probability of this being due

to a different coincidental cause of about 0.001

(White et al. 2012). One follow‐up study

identified the anionic surfactant dioctyl sodium

sulfosuccinate (DOSS), a component of the

dispersant applied during the spill, in floc

removed from one of the corals (White et al.

2014). Another recent laboratory study

experimentally exposed deep‐water corals

(Paramuricea type B3, Callogorgia delta and

Leiopathes glaberrima) to oil, the dispersant

Corexit 9500A and mixtures of the two (Deleo

et al. 2016). Under the conditions of this

experiment the authors report the most severe

declines in health were after exposure to

dispersant alone and the oil‐dispersant mixture.

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The mechanism of the impact from the spill was

not determined, but White et al. (2012)

suggested a possible link to the subsurface

hydrocarbon plume that was detected by

fluorescence and predicted from deep current

models. The principal origins of the plume

remain controversial and the divergent origins

are discussed in most publications dealing with

subsurface hydrocarbon dispersion (e.g. Hazen

et. al. 2010, Camilli et al. 2010, Ryerson et al.

2012, Reddy et al. 2012). In a follow up study,

along with a healthy colony. Hydroid

colonization occurred on some branches

originally covered with the oil‐containing floc

and the probability of hydroid colonization was

directly proportional to the original level of

impact to the colony (Hsing et al. 2013). These

authors emphasize the patchy nature of the

impact on both the community and individual

colony scale and suggest that the impacting

agent may not have been evenly dispersed in

the water at the time of impact.

a b

Figure 4. a) Impacted Paramuricea sp. gorgonian with symbiotic brittle star Asteroschema sp. discovered in

lease block MC294 at a depth of 1,370 m, 11 km southwest of the Macondo well site on November 2, 2010.

Brown floc covering coral tissue contained hopanoid petroleum biomarkers consistent with oil from the

Macondo well. b) Healthy Paramuricea sp. observed in October 2010 at a depth of 360 m over 450 km away

from the spill site. (Images courtesy of Lophelia II, NOAA‐OER/BOEM).

Hsing et al. (2013) found that the median level Fisher et al. (2014a) reported the discovery of

of impact to coral colonies decreased over the two additional coral sites impacted by the

next 17 months as some areas originally DWH spill. One community located 6 km from

covered with floc appeared to recover. One of the Macondo wellhead (MC297) was heavily

the impacted Paramuricia gorgonians impacted, with 49 of 68 corals showing impact

discovered 6 km southeast of the Macondo well to over 5% of colonies in November 2011.

at a depth of 1,370 m is depicted in Figure 4 Another site surveyed at that time, 22 km

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  

distant from the Macondo wellhead (MC344)

and in much deeper water (1,850 m), was more

lightly impacted with only 7 of 30 colonies

showing some evidence of impact from the

blowout. However, the depth of this site

suggests that either the deep‐water plume

impacted the seafloor at greater depths than

either models or the empirical evidence

suggested, or that the impacting agent may

have been oil or dispersant containing marine

snow originating from the sea surface. Data on

the oil content in sediment cores taken from

these sites, as well as studies of the meiofauna

and macrofauna present in the cores, are

consistent with the coral data showing impacts

to these sites resulting from the DWH blowout

(Fisher et al. 2014b).

Additional studies conducted by NRDA

focused on the health and condition of black

corals and octocorals on mesophotic reeftop

biotopes between 65‐75 m depth, in relation to

the number of days below the oil slick and

evidence of exposure to PAHs. Pathologies in

more than 400 octocoral and antipatharian

corals were quantified using methods

consistent with deep‐water studies (Silva et al.

2015). The researchers found elevated PAHs in

coral tissues and sediments, as well as

demonstrable evidence of injury to corals

located below the oil slick at higher frequencies

than ever reported (Silva et al. 2015).

Another study with a Before‐After‐Control‐

Impact research design using video taken since

1985 showed a significant decline in octocoral

health and condition at Alabama Alps Reef,

Roughtongue Reef and Yellowtail Reef,

positioned directly below the oil slick (Etnoyer

et al. 2016). Less than 10% of colonies exhibited

injury before the oil spill, while nearly 50% of

colonies exhibited injury after the oil spill, with

no significant corresponding change at

reference sites (Etnoyer et al. 2016). Odds of

encountering injury after the spill were up to 10

times higher compared to odds of encountering

injured octocorals before the spill. Although

proportions could not be determined, some

injuries were attributed to visible interactions

with fishing line or sedimentation, while others

were attributed to the effects of surface oiling

and dispersants (Silva et al. 2015, Etnoyer et al.

2016). These mesophotic studies extend the

potential footprint of the DWH impacts to sites

nearly 100 km distant from the wellhead.

The NRDA program, and subsequently the

GoM Research Initiative, supported several

years of research into mesophotic and deep

coral habitats to document any potential

injuries to coral associated with the spill. The

principal cruises that contributed information

to and subsequent work on these habitats are

listed in Table 2.

II.4. New Information on the

Distribution of Deep-Sea Corals

Research since 2007 has expanded knowledge

that mesophotic reefs and deep coral habitats

are widespread throughout the GoM (although

generally restricted to relatively rare hard

substrates). Maps showing the locations of

deep‐sea coral records in the GoM are

presented in the Appendix. Several new large

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  

aggregations of corals and sponges were

documented between depths of 50‐2,000 m.

New information has come to light showing

changes with species assemblages on fine depth

scales, over 10s of meters in the mesophotic

zone and over 100s of meters in the deep‐sea.

There is also increased awareness about the

mesophotic diversity of octocorals and black

corals, which has been documented before

(Rezak et al. 1986), but developed further using

molecular techniques. Scleractinian corals have

been shown to occur in zooxanthellate,

azooxanthellate and apozooxanthellate forms,

with the capacity to switch back and forth

between forms (reviewed by Kahng et al. 2010).

Newly identified species in the GoM include

Oculina varicosa aggregations near the Twin

Ridges, south of Apalachicola, Florida (Barnette

2006) and a number of Paragorgia bubblegum

corals when few, if any had been known from

the Gulf before. In deeper waters (300‐1,000 m),

L. pertusa, Leiopathes spp. and Madrepora spp.

have been found to be locally abundant.

Primnoid octocorals (Callogorgia spp.) are also

common in this depth range (Quattrini et al.

2013) and sometimes have catshark egg cases.

Many new observations were made of

Paramuricea octocorals, precious corals like

Corallium, gold corals (family Chrysogorgiidae)

and bamboo corals (family Isididae), which

may occur in abundance as deep as 1,800 m or

more (Doughty et al. 2014).

Habitat suitability models have added new

insight to species distributions, and refined

understanding of how depth and substrate

have a significant influence on deep coral

distributions (Davies et al. 2010, Kinlan et al.

2013, Georgian et al. 2014). New models of

surface productivity, particle flux and bottom

currents are available as well. One previously

established pattern that is well reinforced is that

large, structure‐forming deep corals generally

prefer hard bottom substrates with moderate to

high relief. In the GoM, these substrates can

take the form of banks and mounds, or

authigenic carbonates near hydrocarbon seeps,

or artificial substrates like shipwrecks, as well

as oil and gas platforms. At the time of the last

report (Brooke and Schroeder 2007), the wreck

of oil tanker Gulfpenn was considered one of the

largest aggregations of L. pertusa in the GoM

(Church et al. 2007). Its sister ship, The Gulfoil,

was discovered during Lophelia II in 2010 and

this supported a similar luxuriant community

of L. pertusa (Figure 5). Yet, research since that

time has found even larger aggregations of L.

pertusa in natural environments, including the

largest L. pertusa reef known to date, called

Robert’s Reef.

II.5. Detection of Hard-Bottom

Habitats using Seismic Anomalies

New coral habitats are being discovered

routinely in the GoM, because new types of

information are improving researchers’ abilities

to detect and predict deep coral habitat. With

the continued refinement of industry‐acquired

3D seismic seabed anomaly locations and new

exploration (ground‐truthing) of these

anomalies using ROVs and autonomous

underwater vehicles (AUVs), it is clear now that

there are many thousands of additional habitats

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

with high probability for the presence of corals.

The total number of positive seismic reflectivity

anomalies at the seabed (representing hard

substrate within about 9 m of the seafloor) is

now in the vicinity of 23,000 for the northern

GoM (Figure 6). These discontinuous features

range in size from hundreds of meters in

diameter to many kilometers. However, it

cannot be assumed that all of these seafloor

anomalies correlate to coral and sponge

communities or even that a hard substrate is

exposed on the seabed. The presence of living

of science expeditions for all the

BOEM/NOAA/USGS studies between 2002‐

2012, including many sites chosen for the 2011

and 2012 NOAA Okeanos Explorer missions, and

sites explored around the Macondo well during

the NRDA effort (Fisher et al. 2014a).

II.5.i – Search for Lophelia Reefs using

seismic anomalies

Seabed acoustic anomaly data has been relied

upon heavily in recent years, directing

numerous missions towards the location of

Figure 5. Extensive Lophelia growth on shipwreck Gulfoil at 534 m. This ship sunk in 1942 resulting in 56

years of colonization when first surveyed in 2008. (Image courtesy of Lophelia II, NOAA‐OER/BOEM).

coral or dense hardground communities on

seafloor anomalies needs verification with

visual exploration. Less than 100 sites (less than

0.5% of total) have visual confirmation of

exposed carbonate primarily colonized by deep

corals (although the success rate of finding

living coral habitats is very high as a proportion

of selected anomaly targets). The seismic

targeting process has been a foundational part

deep coral habitat. One example can be

illustrated through the story of a Lophelia

location surveyed in 1955 by Moore and Bullis

(1960). The position of the trawl catch with 300

pounds of Lophelia was reported to the nearest

minute (29° 5’ N, 88° 19’ W), but there were no

features in this area that could support any

substantial Lophelia habitat. The depth of the

collection in the publication correlates almost

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

Figure 6. Continental slope of the northern Gulf of Mexico with a variety of seabed anomalies. The source

3‐D seismic data are extracted for the water bottom and can represent features up to 9 m below the surface.

Red polygons represent interpretation and some consolidation of complex positive reflectivity features.

Other features include pockmarks from gas expulsions and negative reflectivity (potential seeps). (Image

courtesy of BOEM analysis of industry‐acquired 3‐D seismic data).

exactly with seabed 3D seismic anomaly data

(and bathymetry) of a prominent feature

located about 4 miles to the west of the

discovery (or rediscovery) of lush L. pertusa

sites at Viosca Knoll 906 (Schroeder 2007).

Seismic anomalies are publicly available on a

website hosted by BOEM4

. GIS shapefiles for

more than 32,000 anomalies are now available.

4http://www.boem.gov/Oil‐and‐Gas‐Energy‐Program/Mapping‐

and‐Data/Map‐Gallery/Seismic‐Water‐Bottom‐Anomalies‐Map‐

Gallery.aspx

These data have been used to facilitate field

explorations, as well as state‐of‐the‐art habitat

suitability models (Kinlan et al. 2013, Georgian

et al. 2014). The most recent addition to the

seismic anomaly collection is a new category for

Cretaceous‐aged platform‐derived carbonate

along the Florida Escarpment posted in 2012.

One category includes talus detected on the

lower Florida Escarpment. Future explorations

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

Figure 7. Newly discovered expansive Lophelia community at Viosca Knoll lease block 906 (VK906) at a depth

of 390 m. (Image courtesy of Lophelia II, NOAA‐OER/BOEM).

off West Florida can access this information to

guide their surveys.

One of the remarkable deep coral discoveries in

the GoM since 2007 was a discovery in 2009 of a

spectacular accumulation of L. pertusa and

abundant black corals (Leiopathes glaberrima) at

Robert’s Reef. The location of the mound was

associated with a negative seismic anomaly

(Lunden et al. 2013), a short distance from

earlier surveys in lease block VK862 and

adjacent to the same large area of seabed

anomalies associated with most of lease blocks

VK862 and VK906. The Robert’s Reef site

showed that L. pertusa can form large mounds

in the northern GoM (Figure 7), not just thickets

of corals. Age‐dating of piston cores from

Robert’s Reef reaching a depth of 16 m showed

the presence of Lophelia coral throughout the

core dated by a variety of methods to span

approximately 300,000 years (Brooks et al.

2016). This is important because the smaller

thickets known to researchers before 2009

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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Figure 8. Extensive Madrepora oculata with its pink and white skeleton, and other species of corals including

Enallopsammia rostrata and Solenosmilia variabilis from the Green Canyon 852 site at a depth of 1,410 m.

(Image courtesy of Chemo II, BOEM/NOAA‐OER).

suggested the Gulf may be suboptimal habitat

for L. pertusa compared to the North Atlantic.

II.5.ii – Discovery of Madrepora Reefs from

seismic anomalies

Another remarkable discovery since 2007 is that

the branching stony coral Madrepora oculata is

the dominant framework‐forming scleractinian

coral at depths below 800 m (Figure 8). This is

in contrast to shallower depths where M. oculata

occurs primarily as solitary colonies (Brooke

and Schroeder 2007). In these deeper sites at

GC852, Madrepora colonies were found at 1,410

m depth, in a framework with other structure‐

forming corals including Enallopsammia

profunda and Solenosmilia variabilis.

A second Madrepora site occurs at a depth of 886

m in the Mississippi Canyon (lease block

MC118). This site was explored during the

Lophelia II project, and hosts numerous large

thickets. A third site at 1,050 m depth in lease

block AT357 hosts the largest deep‐water coral

assemblage currently known in the central

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

Figure 9. Dense colony of Madrepora oculata with commensal squat lobsters at a depth of 1,050 m in

Atwater Valley lease block 357 (AT357). (Image courtesy of Lophelia II, NOAA‐OER/BOEM).

GoM including Madrepora oculata (Figure 9) and

thousands of colonies of large gorgonian

octocorals Paramuricea sp. with numerous

associated species of epifauna.

II.6. Artificial Hard-Bottom

Substrates; Stepping-Stones for

Lophelia corals?

New information since 2007 suggests that

artificial substrates like shipwrecks and energy

platforms for offshore oil and gas may

5

represent important habitat and are potentially

stepping‐stones that connect populations of

some deep corals (Brooks et al. 2016). To date,

533 structures are known from depths ≥ 50 m

and 53 structures at depths ≥ 300 m. L. pertusa

was first recognized on the deep‐water

Pompano platform at a depth of 400 m during a

NOAA OER cruise in 20035

http://oceanexplorer.noaa.gov/explorations/03bio/welcome.html

(Figure 10). The

Lophelia II program subsequently found L.

pertusa on three shipwrecks and all seven

energy installations investigated (Brooks et al.

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Figure 10. Numerous colonies of Lophelia coral on a portion of the Pompano platform observed in 2003,

representing maximum potential development of nine years since installation in 1994 at a depth of

approximately 305 m. (Image courtesy of John Reed, Harbor Branch Oceanographic Institute).

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

2016). Figure 11b is an example of a thriving

Lophelia habitat that developed on a subsea

manifold structure in lease block MC355 at a

depth of 443 m from the installation date of

1991 to the time of observation in 2012. This

study extended the known depth range for L.

pertusa from 201 to 801 m depth, with the

highest densities occurring near 500 m.

Calculated growth rates on platforms were

higher (≤ 3.2 cm/yr) than those measured using

other methods at natural sites (Larcom et al.

2014). No large gorgonian or black corals were

seen on the platforms, although large octocorals

were present on some of the deeper and older

shipwrecks investigated.

A total of seven wreck sites were investigated

as part of Lophelia II. One site, the tanker

Gulfpenn was once regarded as one of the

largest aggregations of L. pertusa in the GoM

(Church et al. 2007, Figure 11a). The Gulfpenn

survey was repeated from Church et al. (2007),

but several new wrecks were documented in

detail for the first time. Spectacular Lophelia

coral communities were also discovered on the

Gulfoil shipwreck, which were only distantly

seen by Church et al. (2007), and another

shipwreck that was named the Ewing Bank

wreck.

II.7. Detection of Lophelia Reefs

off Western Florida using

Multibeam Data

A large extent of the west Florida shelf and

slope has been mapped since 2007 using high‐

resolution multibeam echosounders aboard

NOAA ships Nancy Foster (Naar 2010) and

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Figure 11. Thriving Lophelia and other associated species on subsea manifold structure in lease block

Mississippi Canyon 355 (MC355) at a depth of 443 m. (Image courtesy of Lophelia II, NOAA‐OER/BOEM).

Museum, Newport News, Virginia). b) Example of Lophelia coral development on the wreck. Photomosaic

depicts numerous colonies nearly completely obscuring the upper bow with some ship structure visible in the

upper right (Image courtesy of Lophelia II, NOAA‐OER/BOEM and S. Lessard‐Pilon).

                                   

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

Figure 11. a) The Gulfpenn, an oil tanker sunk by a German U‐Boat in 1942 (Image courtesy of Mariner’s

Okeanos Explorer6, as well as the Schmidt Ocean

Institute’s R/V Falkor7

. The German expedition

on the R/V Maria S. Merian in 2012 also mapped

the area. The new maps revealed hundreds of

mounds and ridges, some of which were

subsequently surveyed using submersibles and

ROVs. Maps and data from these expeditions

revealed a long continuous escarpment near

400 m depth, many mound features with 10 m

relief or more, and a few isolated ridges with

high vertical relief. Many features are within

the Lophelia depth range (300‐800 m).

6http://oceanexplorer.noaa.gov/okeanos/explorations/ex1202/welc

ome.html

7 http://www.schmidtocean.org/story/show/906

The first reports of Lophelia aggregations on the

west Florida slope were published by Reed et

al. (2006) based on observations from the

Johnson Sea Link submersible. Extensive hard

bottom ecosystems were documented on the

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  

west Florida slope during several research

cruises between 2008 and 2014. These included

two European cruises that conducted seafloor

mapping, water column profiling and

documented L. Pertusa and other corals using

underwater vehicles (Hübscher et al. 2010,

Hebbeln et. al. 2012). Demopoulos et al. (in

revision) describe the physical and

oceanographic characteristics of these deep

reefs and provide estimates of the

accumulations from six cruises (2008‐2012). The

surveys identified mounds and ridges in < 525

m depths capped with living coral colonies,

dominated by L. pertusa. A rocky scarp, running

north‐south for at least 229 km, supported

lower abundances of scleractinian corals than

the mounds and ridges, despite an abundance

of settlement substrata. Several of these Lophelia

aggregations were recommended for protection

to the Gulf of Mexico Fishery Management

Council (GMFMC) (Reed and Farrington 2014).

II.8. Advances in Understanding

Population Connectivity

Although the advancing field of genetics

applies to a wide spectrum of subjects related to

deep corals, it fits well within the discussion of

distribution. Recent advances have been largely

due to the exponential growth in capabilities to

perform genetic analyses. In the deep GoM,

genetic techniques have been used to identify

new coral species (Cairns et al., this volume)

and provide estimates of population

connectivity for the scleractinian coral L. pertusa

(Morrison et al. 2011, Morrison et al., this

volume), the black coral Leiopathes glabberima

(Ruiz‐Ramos et al. 2015) and the octocoral

Callogorgia delta (Quattrini et al. 2015).

Consideration of connectivity among coral

habitats is fundamental to the understanding of

basic ecology, and also critical for consideration

of the sensitivity of populations with respect to

anthropogenic impacts and potentials for

recolonization. Predictions of larval dispersal

for deep corals are difficult due to a lack of

information on timing of reproduction, deep‐

water currents, larval longevity and behavior.

Research since 2007 has shown that L. pertusa

occurs from the deep Garden Banks in the

western Gulf to the deep Pulley Ridge in the

eastern Gulf. Morrison et al. (2011) investigated

the population genetics of North Atlantic and

GoM L pertusa. That paper presented patterns

of connectivity among L. pertusa populations at

various scales and across a large portion of the

species’ Atlantic range from the GoM to

Norway. Using a modeling approach, Morrison

et al. (2011) found four distinct genetic

groupings corresponding to ocean regions:

GoM, coastal southeastern U.S., New England

Seamounts, and eastern North Atlantic Ocean.

In some regions, including the GoM,

connectivity across larger geographic distances

suggests that some larvae are broadly

dispersed. Gene flow within the GoM indicates

high connectivity. Thus, there is high potential

for larvae to repopulate new areas following

impact. However, Morrison et al. (2011)

reported gene flow between ocean regions to be

restricted, indicating more isolation and

concluding that the most effective management

approach for L. pertusa would include regional

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  

conservation networks. L. pertusa in the GoM

were also evaluated for effects from

manipulation, acidification, warming and

deoxygenation with implications of population

genetic variability being responsible for

variable responses (Lunden et al. 2014).

Additional information can be found in the

connectivity spotlight article in this report

(Morrison et al., this volume).

II.9. Distribution of Gorgonian and

Black Coral Communities

Since 2007, black corals and gorgonians are

increasingly recognized as key species within

deep coral ecosystems that provide structure

and habitat, much like branching stony corals

(Etnoyer and Warrenchuk 2007). A

comprehensive study of species diversity across

the entire GoM was published by the Harte

Research Institute (Felder and Camp 2009),

along with a public database curated by more

than 140 taxonomic experts from 15 countries

around the world, titled Biodiversity of the Gulf

of Mexico. Data from this study indicates that

the combined species richness of octocorals and

black corals is higher than scleractinian richness

at all but the shallowest depth in the GoM

(Figure 12). Another specimen‐based study

drawn from museum collections showed that

octocoral diversity peaks at depths between 50

and 800 m and the assemblage varies over a

broad vertical range down to 2,500 m (Etnoyer

2009, Figures 13 and 14), indicating that depth

gradients contribute to patterns of biodiversity

for deep corals in the GoM.

Other field studies since 2007 indicate that

Leiopathes black corals, Callogorgia and

Paramuricea octocorals are widespread in waters

deeper than 200 m. Antipathes, Cirrhipathes, and

Tenacetipathes are among the most commonly

reported black corals from the mesophotic zone

(50‐200 m), whereas Nicella, Ellisella,

Hypnogorgia, Thesea, Scleracis, Bebryce, Swiftia

and Placogorgia are common gorgonian genera

found in depths between 50‐200 m (NOAA

2012).

These corals can occur in high densities,

sometimes over large extents. Previous research

has shown that both black corals and octocorals

provide structure and refuge for demersal

fishes in the mesophotic zone (Weaver et al.

2002). New information is available regarding

the biogeography and vertical distribution of

octocorals in the GoM. Etnoyer (2009)

combined records from national and regional

museums with field samples and cruise reports

to identify hotspots of biodiversity. The study

compared rates of species accumulation for

octocorals in five depth zones: 0‐50 m, 51‐200

m, 201‐800 m, 801‐1600 m and 1600‐3800 m. The

highest rates of species accumulation were

between 50‐200 m and 200‐800 m. Highest

average diversity (Simpson’s D) was in the 50‐

200 m depth range. The study identified some

dissimilarity in the octocoral species

assemblage between the northern GoM and the

Florida platform (Etnoyer 2009). The findings

contrast the hypothesis that deep coral

assemblages are homogenous among the

subregions of the Gulf.

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0

10

20

30

40

50

60

70

80

90

100

0‐5 m 6 ‐ 60 m 61‐200 m 201‐1000 m 1001‐3000 m

Stylasteridae Scleractinia Octocorallia Antipatharia

Figure 12. Coral species richness by taxa and depth strata showing relatively high richness of deep‐water taxa.

(Graph courtesy of Peter Etnoyer and Fabio Moretzsohn using the Biodiversity of the Gulf of Mexico database

at the Harte Research Institute).

Quattrini et al. (2014)’s study, based on samples

collected during Lophelia II expeditions,

supported the idea of two biogeographic regions

in the GoM using measures of phylogenetic

diversity, and extended this concept into deeper

waters. Highest species richness for octocorals

occurred at the shallowest sites (< 325 m) and the

deepest sites surveyed (1,800‐2,500 m).

Specimens were collected from 31 hard bottom

sites in the northern and eastern GoM between

2008‐2011. This pattern was different from that

of museum samples, and opposite of the typical

pattern of deep‐sea diversity in the GoM, which

is expected to peak at mid‐slope depths

(Gallaway et al. 1988, Rowe and Kennicutt 2009).

The research described transitions in octocoral

assemblages at depth breaks near 325, 425, 600,

1,100 and 2,100 m. Octocoral assemblages on

the west Florida slope represented a separate

biogeographic group within the 425‐600 m

depth range compared to other groups located

in the north central GoM. Using molecular

tools, these authors also report on 12 species

never before collected in the GoM, including

three that are likely new to science.

Species within the genus Callogorgia are the

most common and abundant gorgonian

octocoral taxa reported from the upper

continental slope (200‐1,000 m) in the GoM

(NOAA 2015). Quattrini et al. (2013)

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Figure 13. Analysis of gorgonian species diversity by depth range from Gulf of Mexico museum collections

showing high diversity in the 50‐200 m depth range. The number of zones on the species accumulation curves

correspond to regional zones defined in Etnoyer (2009). (Graphic reprinted from Etnoyer 2009.)

of species accumulation were between 50 200

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  

‐

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  

                

                                 

               

Figure 14. Analysis of the vertical distribution of Gulf of Mexico gorgonian families and genera, as derived

from museum collections. (Graphic reprinted from Etnoyer 2009).

m depth and 200 800 m depth. Highest average

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

documented strong niche segregation with

depth in the genus using specimen collections,

video data, environmental niche modeling and

DNA barcoding. C. gracilis occurred at the

shallowest sites (depths up to 320 m), C.

americana was found in mid‐depths (339‐384 m)

and C. delta was found at the deepest depths

(403‐914 m) surveyed. C. delta was also

associated with cold seeps, indicating that this

species may have adaptations to living in areas

of increased hydrocarbon seepage.

Following this study, Quattrini et al. (2015)

examined whether depth could serve as an

isolating factor in populations of C. delta.

Significant genetic differentiation occurred

across seven sites spanning 400 km of distance

and 400 m of depth. Greater genetic

differentiation was observed with vertical

distance, rather than geographical distance;

however, geographical distance may also play a

role in limiting gene flow. The study concluded

that water mass boundaries isolate populations

across depth, subsequently leading to adaptive

divergence with depth (Quattrini et al. 2015).

Another recent study examined the population

structure of the black coral L. glaberrima in the

deep Gulf (Ruiz‐Ramos et al. 2015). These

authors found that although different color

morphotypes of this coral could not be

distinguished genetically, they did show

distinct differences in distribution. The study

also found evidence of two distinct populations

of L. glaberrima, one of which showed evidence

of the ability for long range dispersal (and was

present at all sites examined) and another

which was restricted to VK826 and VK906

where it occurred sympatrically with the other

lineage. These results indicate that coupled

ecological and evolutionary processes are

important in structuring the distribution of

deep corals in the GoM.

Doughty et al. (2014) investigated the

occurrence, density and size distributions of

one of the most common gorgonian octocorals

in the Gulf below 200 m (Paramuricea spp.).

Similar to the results of Quattrini et al. (2015),

the species of Paramuricea present in the GoM

(based on molecular haplotypes) were partially

segregated by depth. The authors concluded

that as a result of their sparse distribution, low

recruitment rates and slow growth rates, these

deep‐sea gorgonian octocorals are highly

susceptible to anthropogenic threats.

NOAA’s Deep Sea Coral Research and

Technology Program (DSCRTP) has developed

a national database of deep‐sea corals and

sponges with nearly 300,000 coral records

including black corals and octocorals. This has

greatly broadened the number of records of

deep coral locations (see annex to this chapter).

Habitat suitability models were developed from

these occurrences to help fill gaps in the survey

extent and to increase the reliability of

predictions of coral locations that have never

been surveyed before (Kinlan et al. 2013).

Habitat suitability models have contributed

immensely to a better understanding of the true

extent of potential deep coral habitats at broad

regional scales, as well as the mesoscale

(Georgian et al. 2014).

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II.10. Continental Shelf Mesophotic

Habitats

II.10.i – Northwestern GoM

An updated biological habitat characterization

scheme was released for the reefs and banks on

the outer continental shelf of the northwestern

GoM, including FGBNMS (Hickerson et al.

2008, Schmahl et al. 2008), based on the results

of sampling efforts since 2003. Nuttall (2013)

compared field identifications of black coral

species (collected from the FGBNMS and other

historical data) with laboratory identifications,

and showed that species‐level field

identifications are not reliable and genus level

identifications are more accurate. Nuttall (2013)

also contains habitat suitability maps for

antipatharians. Wicksten et al. (2014) described

crustaceans associated with black corals,

including a new species of crustacean, the squat

lobster Uroptychus marissae (Baba and Wicksten

2015). Rodriguez (2015) studied black corals on

nearby South Texas Banks showing highest

abundance and diversity of black corals on the

tops of banks compared to the flanks. An

isolated observation of the antipatharian

Plumapathes pennacea was reported by Boland

and Sammarco (2005) on the crest of the East

Flower Garden Bank at a depth of 22 m.

Opresko et al. (in press) provided a guide to the

antipatharians of the FGBNMS, which is useful

throughout the region.

On the Flower Garden Banks, octocorals are

absent shallower than 50 m (Bright et al. 1984),

but a total of 24 species were documented in 16

genera (Etnoyer 2009) in deeper waters (52‐130

m). The octocoral assemblage was stratified by

depth, with a shallow group (50‐70 m) and a

deeper group (70‐125 m). Hotspots of octocoral

species richness and abundance were identified

at the base of East and West Flower Garden

Banks. Furthermore, four different octocoral

assemblages were recognized at among six

banks within a similar depth range, including

Flower Gardens. Thus, nearby features 20‐40

km apart may have very different species

composition. One surprising result of the study

was a high diversity of octocorals on low relief

hard‐bottom habitats between East and West

Flower Garden Banks, on a feature now

referred to as Horseshoe Bank.

The publications resulting from BOEMs

Potentially Sensitive Biological Features

investigations will provide insights into the

distribution, diversity, dominance and

structure of the mesophotic communities in the

region, as well as the fine‐scale geomorphology

of a set of 14 banks in the northern GoM

(Sammarco et al. 2016). This will also contribute

to our understanding of the relationship

between the fine‐scale geomorphology and

species richness in these habitats at a variety of

spatial scales (Sammarco et al. 2016).

Considerable range extensions of fish

populations have resulted from the

investigations in this region. An effort is

currently underway to organize and publicize

this data.

II.10.ii – Pinnacle Trend Region

Considerable new research occurred since 2010

on the mesophotic reefs at Alabama Alps,

Roughtongue, and Yellowtail in the Pinnacle

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  

Trend region off Louisiana, Alabama and

Mississippi (Figure 3), as well as on Coral Trees

Reefs and Madison Swanson Marine Reserve

off Florida. These studies also found differences

in species composition between the two groups

of reefs at similar depths (Etnoyer et al. 2016).

Pinnacle Trend sites were dominated by the

octocorals Swiftia sp., Thesea nivea and

Hypnogorgia sp., while at the Florida sites,

Thesea rubra and Placogorgia sp. were most

common. These differences in species

composition are consistent with the idea of a

biogeographic boundary for deep corals

between the northwestern and the eastern

GoM. Differences could be driven by

differences in seasonal temperatures, salinity,

productivity or underlying substrate. In

contrast, habitat suitability models developed

for Swiftia, Thesea and Hypnogorgia indicate that

these taxa are broadly distributed (Kinlan et al.

2013), but models can widen distributions

beyond direct observations and rely on ground‐

truthing. Much remains to be explored in terms

of population connectivity and environmental

tolerances for the mesophotic sea fans and

black corals.

II.11. Species Range Extensions

Many species of habitat‐forming deep corals

were encountered in the GoM for the first time

(Figure 15), or in exceptional abundance, since

the last report in 2007. Bubblegum corals

(Paragorgiidae) were thought to be rare in the

GoM, but they have been sampled and

observed numerous times since 2008, at depths

ranging from 360‐1,800 m depth, in both the

northwestern and eastern parts of the Gulf

(Brooks et al. 2016). Precious corals (Corallidae)

and Iridogorgia spiral corals (Chrysogorgiidae)

have also been observed. A list of species

discovered during Lophelia II or Chemo III, but

not included in Cairns and Bayer (2009) is

shown in Table 3. A comprehensive list of deep

coral species in the GoM is provided in an

annex to this chapter. This annex is modified

from Cairns and Bayer (2009) to include

newspecies occurrences and range extensions

since the original publication. Clearly, the more

we explore, the more we discover and observe.

II.12. Occurrence and Distribution

of Deep-Sea Sponges

Limited data still exist on sponges in the deep

GoM despite intense research over the last ten

years. A map showing the locations of deep‐sea

sponge records in the GoM is presented in the

Appendix. The paucity of research on these

important taxa was identified but not fully

addressed in the 2007 volume (Lumsden et al.

2007). Rützler et al. (2009) indicated the

presence of 66 taxa for the deep‐sea from 12

orders with three orders representing the

majority of the total; Lithistida (15),

Poecilosclerida (14) and Hexactinosida (12). No

annotations or references in that summary are

dated after 2004. The last comprehensive study

of the deep GoM by Rowe and Kennicutt (2009)

consisted of widely separated sampling stations

ranging in depth from 200 to 3,750 m spanning

across most of the northern GoM and also

352
                          

       

     

                                   

                                 

                         

Figure 15. Several species and genera of deep‐sea gorgonian octocorals were recorded in the Gulf of Mexico for

the first time since 2007, including species in the genera a) Callogorgia, b) Iridogorgia, c) Corallium, d)

Paragorgia, e) Paramuricea, and f) Keratoisis (see annex list of deep‐sea coral species).

                             

                               

Family Species 

Paragorgiidae Sibogorgia cauliflora*, Paragorgia wahine*, Paragorgia johnsoni, Paragorgia regalis 

Corallidae Hemicorallium niobe (=Corallium niobe), Corallium medea 

Primnoidae Callogorgia delta 

Chrysogorgiidae Iridogorgia splendens, Iridogorgia magnispiralis 

Plexauridae Paramuricea biscaya 

Clavulariidae Clavularia rudis 

Aquaumbridae (new family described in 2012 from the tropical eastern Pacific) 

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

Table 3. Octocoral range extensions documented from the Chemo III and Lophelia II study. (*Confirmed by

taxonomic experts S. Herrera and J. Sanchez. Designations of new species await additional genetic data and

morphological confirmation).

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  

included some stations in Mexican waters.

However, not much was reported on sponges

and results were primarily qualitative.

A prior Gulf‐wide study (Gallaway et al. 1988)

was conducted in the mid‐1980s and represents

a more significant body of data on GoM deep‐

water sponges and also includes one of the few

records of a high‐density sponge area on the

GoM slope. This study included 60 stations that

spanned most of the northern GoM ranging in

depth from 300 to 3,000 m. Gallaway et al.

(1988) report 39 taxa of Porifera with 22

identified to species. Densities of small sponges

from boxcore transects ranged from 0‐137/m2

with the maximum density observed at a single

site at Station E5 during the fourth of five

cruises at a depth of 2,902 m. However, the

species observed were not reported (Gallaway

et al. 1988). The ten most abundant taxa

identified from boxcore samples as part of the

macrofauna category, and mostly small in size

and not structure forming, were only identified

to genus or family. These included the genera

Stelletta, Sycon, Thenea, Stylocordyla, Thenea,

Mycale (species A and B) and Microciona, as well

as the families Suberitidae and Suberotodae.

Sponges were also reported as part of the

quantitative benthic photography component of

Gallaway et al. (1988) using a drift camera

maintained 2 m off the bottom at the same 60

stations. Hard bottom was not targeted.

Densities from imagery exceeded densities

from trawling at all stations. Maximum sponge

density from imagery was 16,514 per hectare at

one station at a depth of 1,400 m. Overall

density of Porifera from benthic photography

was approximately 450 individuals per hectare.

In the mesophotic region, research conducted

by the FGBNMS has collected about 50 sponge

samples in the 50‐150 m depth range in the NW

GoM. These samples were identified with the

help of Dr. Christi Savarese, and Klaus Rützler

at the Smithsonian National Museum of

Natural History. An interesting glass sponge

habitat was identified at Elvers Bank and

warrants further investigation. Results from

these efforts are available in poster form.

8

8http://flowergarden.noaa.gov/document_library/scidocs/spongep

oster.pdf

II.13. New Information of Species

Associations and the Role of

Corals and Sponges as Habitat

Work has been done on deep coral associates

including results from earlier studies (Lophelia

I) ranging from microsymbionts to crustacea

and fish. Wicksten et al. (2014) described

crustaceans associated with black corals, as well

as a new species of crustacean, the squat lobster

Uroptychus marissae (Baba and Wicksten 2015).

Giarad et al. (2016) reported on the benefits

provided by the commensal ophiuroid

Asteroschema clavigerum living on Paramuricea

colonies impacted by the DWH spill. Recovery

from visible impact and hydroid colonization of

Paramuricea colonies was negatively correlated

with distance from the ophiuroid. Branches

within the area of influence of the ophiuroid

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  

were also more likely to recover, indicating

likely benefits from its association with A.

clavigerum through the physical action of

ophiuroids removing material depositing on

polyps. Cordes et al. (2008) reported 68

invertebrate taxa that were identified in close

association with the L. pertusa coral framework

in the northern GoM. Lessard‐Pilon et al. (2010)

report high diversity of fish and invertebrates

in L. pertusa structure and that the highest

diversity is associated with thickets containing

high proportions of dead L. pertusa. Because L.

pertusa in the central and northern GoM are

often associated with current or historical seeps

through colonization of derived carbonates,

Becker et al. (2009) used stable isotopes to

investigate a potential relation between L.

pertusa communities and seep primary

production. With the exception of a single

gastropod (of 33 species tested), the authors

found no evidence for a nutritional dependence

of L. pertusa, or the fauna associated with the

coral colonies, on seep primary production.

Much of the work of the USGS Lophelia II team

is contained in the regional publication list

including Kellogg et al. (2009), Ross et al. (2012)

and Demopoulos et al. (2010). Final reports

from this suite of major studies also contain

significant new information on deep coral

associates, trophic relationships of associated

species groups, substrate infauna and new

results from microsymbiont studies.

II.14. New Research Priorities

Research priorities for deep coral in the GoM

are many. There continues to be a need for

baseline data in preparation for future

catastrophic incidents and potential habitat

impacts due to ecosystem impacts from climate

change. The DWH incident clearly illustrates

this need. Additionally, it is necesssary to

understand long‐term responses to major

environmental events. Below we outline several

research priorities to increase knowledge on

deep‐sea and mesophotic habitats in the region.

• Known deep coral and sponge sites need to

be mapped at high resolution. This is the

first step in exploration and characterization

of these resources.

• Continued exploration and characterization

of known sites in order to manage and

protect resources.

• Long‐term monitoring of deep coral

habitats at a variety of sites throughout the

GoM, including but not limited to, the sites

impacted by the DWH incident.

• Growth rates, damage and aging of deep

corals, as well as the natural abundances of

fauna both directly and indirectly

associated with deep corals.

• Enhance the predictive capability (i.e., use

of remote sensing rather than visual

detection) for identifying the distribution of

deep coral including a modeling approach.

NOAA’s DSCRTP has begun a modeling

project for the GoM. Additional ground‐

truthing and other research could be

required. Ideally this would overlap with

other research priorities.

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• Understand two major components that

determine the success of recolonization of

potentially impacted habitats; 1)

reproduction and development of species of

concern (e.g., Lophelia, Madrepora, Leiopathes

and Paramuricea) and 2) physical factors

determining the transport of reproductive

propogules of species of concern (both

gametes and larvae).

• Investigations of genetic connectivity

between populations are needed to

effectively manage these resources.

• Archiving of invertebrate specimens

including corals and related community

species will continue with an ongoing

contract between BOEM and the

Smithsonian National Museum of Natural

History. Additional scope of this program

will include tissue archiving in their

recently completed biorepository, as well as

potential genetic barcoding. This effort

should be expanded as much as possible

and extended to other agencies involved in

collection of deep‐sea coral specimens.

II.15. Update on Management:

New Information on Stressors

II.15.i – Oil and gas development 

Brooke and Schroeder (2007) reviewed the

stressors on deep coral communities in the

northern GoM. Oil and gas development was

identified as having a perceived medium level

of threat to these communities – the highest

relative threat level for human activities in the

Gulf. As part of continued offshore

development in the GoM, a total of 1,906 wells

were drilled in water depths below 50 m from

January 2007 to August 2015. Of that total, 1,066

were at depths below 300 m. Information on

wells can be accessed through a variety of

query options at a regularly updated internet

site at BOEM.

9 Other than some visible linear

physical impacts from anchor cables reported in

Schroeder (2007), there have been no

documented impacts to deep coral habitats

from routine oil and gas activities.

The DWH oil spill and associated use of oil

dispersants showed how a major accident could

affect deep coral communities. Numerous

additional research publications directly

involving deep corals and the associated

communities will become available over the

next few years as a direct result of research

funded though federally required oil spill

damage assessment, the GoM Research

Initiative, as well as a spectrum of additional

research funding related to the DWH event.

One thing that is clear from NRDA benthic

studies is that deep corals are important

sentinel species that are useful for monitoring

and vulnerable to oil pollution impacts.

II.15.ii – Fishing threats

9https://www.data.boem.gov/Well/Borehole/Default.aspx

Bottom longlines for reef fish are a potential

threat to corals and sponges in shelf‐edge

depths, because these fisheries target the same

complex reef habitats where the target species

of groupers and snapper occur including

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  

snowy grouper, Warsaw grouper and tilefish.

Lost longline gear has been observed

throughout the mesophotic reefs of the

Northwestern GoM. Abandoned fishing lines

have been observed with some frequency near

Coral Trees Reef and Madison Swanson South

Reef (MSSR), sometimes wrapped in and

around gorgonian corals. The reefs are situated

in an offshore region between Pensacola and

Tampa, Florida known as the northern grounds

for the GoM bottom longline fishery and the

area of the Gulf where fishing for reef fish

actually began in the 1800’s (Prytherch 1983).

Intense fishing pressure at MSMR led to closure

in 2000, when the GMFMC prohibited all

bottom contact gear in the reserve. Low levels

of bottom longline fishing effort were reported

from the Pinnacles region in 2001, but the

majority of effort was concentrated in the

eastern GoM, near Tampa, Florida (Scott‐

Denton et al. 2011). Observations of fishing line

on Pinnacle Trend sites were few compared to

Coral Trees Reef and MSSR (Etnoyer et al.

2016). Anchoring associated with specific types

of fishing, such as bandit reel fishing, is also a

threat to mesophotic reef systems.

There is little available information regarding

new fisheries operating deeper than 300 m in

the GoM that could significantly impact deep

corals. An overview of this subject was a

component of the Lophelia II project and is

presented in Brooks et al. (2016). Commercial

fishing pressures on deep coral in the GoM

remain low compared to other regions;

however, discarded fishing line has been

documented in deep coral communities and

may pose a threat to these coral communities

(Fisher et al. 2014a). The only two fisheries

identified as having some potential impact to

deep‐water scleractinian coral habitats, such as

Lophelia and Madrepora, are limited to golden

crab (Chaceon fenneri) and royal red shrimp

(Pleoticus robustus). A fishery for the deep‐sea

golden crab has not developed since its early

description in the literature (Brooks et al. 2016).

However, this pot and trap fishery does come

in contact with the bottom near 500 m depth

where deep corals occur. The pots are strung

together and retrieved by grappling hook

before being dragged to the surface, so there is

some potential for severe bottom disturbance in

sensitive coral habitats. Golden crab is not

currently a managed fishery in the GoM, but

there is a small fleet operating off the

southeastern U.S. coast.

There is some evidence of a golden crab fishery

operating in and around Lophelia aggregations

off the west coast of Florida. Golden crabs, a

discarded crab pot, and broken Lophelia coral

rubble were observed in 2012.

10 The feature has

not been revisited since that time. There are

only eight reports of golden crab landings in

the NMFS fishery logbook databases from 1995

to 2004.

11

10http://www.schmidtocean.org/story/show/915 11http://www.st.nmfs.noaa.gov/st1/commercial/landings/annual_l

andings.html

357

Landings data include small catches,

some likely as by‐catch, ranging from 25‐641

MT over a period of eight years. The last

reported data for golden crab landings in the

GoM from the National Marine Fishery Service
                          

       

     

         

         

                 

         

               

           

           

               

               

               

           

             

                 

                 

                 

                 

               

           

                   

           

                       

                   

           

           

           

               

             

           

     

             

             

               

             

                 

             

           

             

         

         

           

             

                

             

               

            

           

              

             

             

               

               

           

               

           

               

           

             

           

               

                 

         

               

             

              

             

               

               

             

                 

STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

(NMFS) reference site for annual commercial

fishery landings was 2004.

Royal red shrimp are fished in the GoM using

bottom‐contact trawling on the continental

slope. The vessels target soft bottoms, but are

occasionally close to known deep coral

ecosystems including those in VK862/906. This

species represents a small portion of the overall

U.S. shrimp fishery (Stiles et al. 2007). The

shrimp are sometimes observed close to and in

association with deep corals (Ross 2005).

Landings data from the last available NOAA

data records for this species in 2014 is recorded

for the Gulf States of Alabama (59.7 MT). A

small catch was also reported in 2014 for the

west coast of Florida (17.8 MT). The most recent

data from Louisiana was in 2013 (2.8 MT).

Texas reported relatively small royal red

shrimp catches of less than 1 MT only in 2004

and 2012. Alabama landings diminished from

141 MT in 2009 to 102 MT in 2010, but rose to

159 MT in 2011 and 144 MT in 2012, remaining

relatively consistent since the 1990s.

II.15.iii – Invasive species

The invasion of the Indo‐Pacific lionfish (Pterois

volitans and P. miles) also poses a different kind

of threat to GoM deep coral and sponge

habitats. Lesser and Slattery (2011) report loss

of biodiversity and resilience of nursery areas

(mangroves), coral reefs, mesophotic

ecosystems and artificial reefs, as well as

cascading impacts across food webs. Based on

the gut content analysis, and the size of

potential prey, it should be acknowledged that

the lionfish invasion poses a threat to prey in

deep coral habitats, and this may have

cascading effects throughout the lionfish depth

range. One potential impact is that the

reduction of herbivorous fish species

abundance, indirectly impacts corals by

fostering algal growth (Albins and Huxon

2013).

Invasive lionfish have been reported down to

depths of 300 m (Albins and Hixon 2013).

Lionfish were first captured off the northern

Yucatan peninsula in December 2009 and in the

northwestern GoM in September of 2010.

Numbers of sightings have been increasing

exponentially in the northern Gulf since 2011.

Since lionfish were first observed in the

FGBNMS in 2011, 2,614 lionfish sightings have

been reported at depths ranging from 17‐96 m,

inclusive of both the shallow reef cap and

mesophotic habitats through 2015. From this

total, 1,484 (56.8%) have been removed. To date

approximately 600 lionfish have been reported

from other reefs and banks in the northern

GoM, outside of the FGBNMS, primarily

during ROV surveys at depths ranging between

50‐177 m (Nuttall et al. 2014).

While lionfish were first observed by divers on

all three banks of the FGBNMS in 2011, they

were not recorded in long‐term monitoring

surveys until 2013, most likely due to the

limited number of lionfish present during the

first two years (Johnston et al. 2016a). Lionfish

sightings from these surveys, occurring on the

shallowest portion of the reef caps (17–27 m),

varied among the three banks. In 2013, lionfish

sighting frequency at East Flower Garden Bank

was 25%, 33% at the West Flower Garden Bank

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

and 5% at Stetson Bank. Sighting frequency in

2014 doubled at East Flower Garden Bank

(50%) and increased at West Flower Garden

Bank (40%), but decreased at Stetson Bank

(2.9%). In 2015, sighting frequency decreased at

East Flower Garden Bank (17%), increased at

West Flower Garden Bank (60%), and was

below 1% at Stetson Bank (Johnston et al. 2016

a, b). In 2015 long‐term monitoring surveys,

average lionfish density was 0.55 per 100 m2 at

the East and West Flower Garden Banks.

II.16. Management Actions

Much has happened in the GoM since 2007,

particularly with regard to the management of

the energy industry. Some new policies

preceded the DWH oil spill, and it is likely that

more management actions will result from

ongoing reviews and research. Numerous

reforms to the offshore oil and gas industry

were also enacted in response to the DWH

blowout and resulting oil spill in 2010.

II.16.i – Management of oil and gas

development

BOEM manages the exploration and

development of the nationʹs offshore energy

resources. Since 2007, the agency made changes

to two specific regulatory policies for the oil

and gas industry dealing with biological

communities in the GoM, in addition to

numerous changes in regulatory policies

related to drilling safety.

BOEM supplements regulations that govern

energy development operations on the Outer

Continental Shelf through a regulatory

mechanism called Notices to Lessees (NTL12

12http://www.boem.gov/Notices‐to‐Lessees‐and‐Operators

).

For example, NTL 2009‐G40, titled Deepwater

Benthic Communities, became effective in

January 2010 and it increased the distance of

avoidance from sensitive deep‐water biological

communities, including both chemosynthetic

communities and deep coral habitats, for

drilling discharges (610 m) and anchoring (152

m). This effectively doubled the distance of

avoidance from prior NTL regulations. NTL

2009‐G40 applies to all oil and gas activities,

including exploration and production drilling

plans, as well as pipeline applications, in water

deeper than 300 m (the depth was raised from

400 m to encompass newly discovered deep

coral communities). As described in Schroeder

(2007) there was some evidence of physical

impacts to deep corals from anchoring activities

in the past, specifically at the coral communities

in lease block VK826. There have been no

similar impacts observed to deep coral sites

related to energy development activities on

deep corals since 2002, with the exception of the

DWH incident.

A second NTL (NTL 2009‐G39, effective date

January 2010), titled Biologically‐Sensitive

Underwater Features and Areas, applies to

water depths shallower than 300 m and

protection to sensitive biological features. In

addition to previously established protective

measures for topographic features and live

bottom areas including the Pinnacle Trend

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

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  

region, a new avoidance category of bottom

feature was created for this NTL. This new

category, titled potentially sensitive biological

features, is defined as features of moderate to

high relief (about 2 m) that are not protected by

other biological lease stipulations. This

addressed the increasing awareness that

extensive areas of exposed hard bottom and

associated communities (including corals) were

not included in No Activity Zone boundaries of

named topographic features (e.g., Bright Bank,

Geyer Bank and Sonnier Bank) or within the

defined areas of high diversity features in the

Pinnacle Trend. Although this major shift in

protection for continental shelf features was

first implemented in 2005 (part of earlier NTL

2004‐G05), it was not addressed in Brooke and

Schroeder (2007) and is continuing to evolve

with extensive field research efforts in these

habitat areas.

Comprehensive reforms and fundamental

changes were made to offshore oil and gas

regulation and oversight in response to the

DWH explosion and resulting oil spill in the

GoM to maintain responsible oil and gas

drilling and production on the U.S. Outer

Continental Shelf. Beyond the fundamental

reorganization of the former regulatory agency,

Minerals Management Service (MMS) into two

new, independent bureaus – the Bureau of

Safety and Environmental Enforcement (BSEE)

and BOEM, numerous regulatory policies have

also been enacted since 2010. With

consideration of the impact to deep corals, the

following new policies have a direct influence

on the increased safety of deep‐water drilling

and future accident events. Other new policies

such as more exacting inspection criteria and

heightened standards for well design, casing

and cementing are also indirectly tied to the

issue but will not be detailed here.

1) One NTL was implemented shortly after the

DWH spill, NTL 2010‐N06 now updated to

NTL 2015‐N01, “Information Requirements

for Exploration Plans, Development and

Production Plans, and Development

Operations Coordination Documents on the

OCS” requires operators to demonstrate

they are prepared to deal with the potential

for a blowout and worst‐case discharge.

2) A drilling safety rule implemented in 2010

makes mandatory several requirements for

the drilling process that enhance safety of

oil and gas drilling operations on the Outer

Continental Shelf. It addresses both well

bore integrity and well control equipment

and procedures requiring two independent

test barriers across each flow path during

well completion activities. BSEE inspectors

must now be on location and observe the

Blow Out Preventer (BOP) testing prior to

drilling commencing at the rig site.

3) Before receiving approval for deep‐water

operations, all operators now must

demonstrate the capability to contain a

subsea blowout like the one seen in the

DWH spill. Operators demonstrate that

they have access to all necessary equipment

for subsea well control and containment,

including a capping stack. As a result, there

is now containment equipment available for

industry deployment.

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  

The most recent five‐year leasing plan for GoM

energy development (2017‐2022) is described on

the BOEM website. The five‐year program

consists of a schedule of oil and gas lease sales

indicating the size, timing and location of

proposed leasing activity that will best meet

national energy needs for the five‐year period

following its approval. A total of ten separate

lease sales have been proposed that will include

the entire available GoM leases as opposed to

separate sales for the Central, Western and

Eastern Planning areas. The total submerged

lands incorporated in the next five‐year cycle

include an area of approximately 647,499 km2

(160 million acres).

Potential future management actions include

the consideration of expanded No Activity

Zones for the more formal inclusion of densely

spaced habitat features adjacent to existing

banks (potentially sensitive biological features).

II.16.ii – Other management actions

In June, 2016, the FGBNMS released a draft

environmental impact statement outlining a

range of alternatives for boundary expansion,

including five alternatives ranging from no

action (current sanctuary area of 56.21 miles2) to

a comprehensive plan to protect known high

value benthic habitats and cultural resources in

the north‐central GoM. A preferred alternative

was identified, which proposes an expansion to

383.19 miles2, and includes Horseshoe (between

East and West Flower Garden Banks), MacNeil,

Rankin, 28 Fathom, Bright, Geyer, Elvers,

Sonnier, McGrail, Rezak, Sidner, Bryant,

Bouma, Parker and Alderdice Banks. All of

these banks harbor significant habitats that

include deep corals and sponges. In addition to

these proposed expansion sites, the document

also proposes to amend the boundaries of the

three banks currently in the sanctuary (East and

West Flower Garden and Stetson), to better

encompass and therefore protect the deep coral

habitats. After a period of public scoping, a

Final Environmental Impact Statement will be

developed and likely released in 2018.

NOAA’s NMFS and the GMFMC have

authority over fisheries in federal waters of the

GoM. There have been no major changes in

federal fishing regulations affecting deep coral

areas since essential fish habitat designations in

2006. In June 2015, the GMFMC considered

recommendations from its coral advisory

bodies to evaluate 47 sites for Habitat Areas of

Particular Concern designations and to protect

them from fishing gear impacts (DSCRTP 2016).

These sites are distributed throughout the GoM

from Texas to Florida, and all have confirmed

presence of multiple deep‐sea coral taxa

including the South Texas Banks, shelf and

slope sites in the northern GoM and west

Florida Platform including sites in Mississippi

Canyon, Green Canyon, Garden Banks,

Pinnacle trend and Viosca Knoll. The GMFMC

plans to seek input from the fishing community

on this proposal before officially starting a

process to amend the Fishery Management Plan

for Coral and Coral Reefs with new Habitat

Areas of Particular Concern designations.

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  

II.17. New Management Priorities

Following on from the ongoing Potentially

Sensitive Biological Features shelf‐edge study

described above, additional high‐resolution

mapping will be needed for GoM topographic

features where only low‐resolution bathymetry

is known (most from early 1970s). Additional

mapping will lead to further investigations of

any relevant benthic features and consideration

of additional management protective measures.

Results from this study will allow refinement of

avoidance of energy development activities

from benthic features occurring outside of No

Activity Zones established by BOEM, as well as

help guide development of other potential

protective measures by other agencies.

If future oil and gas leasing moves farther east

in the GoM, additional investigations on the

west Florida shelf and slope will become a

higher priority to inform management

decisions for protective measures within this

area that is much different than authigenic

carbonate features of the rest of the GoM.

Newly released BOEM shape files of water

bottom seismic anomalies in this area will be

very informative in the future, by allowing

targeted research. Additionally, these files will

be fundamental for management decision‐

making related to avoidance criteria of hard

substrate depicted in survey databases.

III. Conclusion

An extensive amount of research on deep corals

has been performed in the GoM since 2007,

both prior to the DWH incident and afterwards,

leading to increased awareness among

managers and the public. This awareness has

resulted in significant management actions to

protect deep coral habitats in the mesophotic

zone and deeper waters. Even though much

less targeted research has been devoted to

deep‐water sponge ecosystems, it appears

sponge communities (at least high‐density

communities) are relatively rare in the GoM

compared to corals. The DWH incident was a

landmark event with major revelations related

to deep spill impacts including the unclear

dynamics of deep plumes, extensive subsurface

and surface dispersant use, the potential for

impacts to deep‐water habitats from horizontal

transport of hydrocarbon plumes and from

oiled marine snow. Increased understanding of

genetic diversity, age, growth, reproduction

and distribution of deep corals will greatly

benefit management efforts, but much remains

to be discovered and described. The availability

and interpretation of GoM seismic reflectivity

data have greatly facilitated discovery of new

deep coral habitats throughout the northern

GoM. The addition of more sophisticated

modeling and ground‐truthing efforts has

greatly enhanced our understanding of deep

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  

coral distribution in the GoM. New multibeam

data and exploration efforts along the West

Florida Shelf has led to the subsequent

discovery of large aggregations of sensitive

coral habitats, and these should be investigated

in the future to determine their extent and

degree of vulnerability to climate change and

other anthropogenic impacts. There continues

to be a need to map, explore and characterize

vast areas of the GoM and to better understand

the biology and ecology of deep corals and

sponges in order to provide the understanding

necessary for the development of appropriate

regulatory and protective measures.

IV. Acknowledgements

We thank Sandra Brooke, Cynthia Cooksey,

Andrea Quattrini and Paul Sammarco for their

thoughtful reviews and expert comments on an

earlier draft of this chapter. Their contributions

and in‐depth suggestions greatly enhanced the

quality of the final product. Reviewers in

BOEM included Brad Blythe, Rodney Cluck

and William Brown. We also thank Robert

McGuinn for generating maps. Tom Hourigan

provided helpful guidance through the lengthy

period of early writing and development. We

thank Daniel Wagner for his assistance in final

formatting and layout of this chapter.

Online Annex – Comprehensive list of deep‐

sea corals in the U.S. Gulf of Mexico region link:

https://deepseacoraldata.noaa.gov/library/2017‐

state‐of‐deep‐sea‐corals‐report

V. Literature Cited

Albins MA, Hixon MA (2013) Worst case scenario:

potential long‐term effects of invasive predatory

lionfish (Pterois volitans) on Atlantic and

Caribbean coral‐reef communities.

Environmental Biology of Fishes 96:1151‐1157

Baba K, Wicksten M (2015) Uroptychus minutus

Benedict, 1902 and a closely related new species

(Crustacea: Anomura: Chirostylidae) from the

western Atlantic Ocean. Zootaxa 3957:215‐225

Barnette MC (2006) Observations of the deep‐water

coral Oculina varicosa in the Gulf of Mexico.

NOAA Technical Memorandum NMFS‐SEFSC‐

535

Becker EL, Cordes EE, Macko SA, Fisher CR (2009)

Importance of seep primary production to

Lophelia pertusa and associated fauna in the Gulf

of Mexico. Deep‐Sea Research Part I:

Oceanographic Research Papers 56:786‐800

Boland GS, Sammarco PW (2005) Observations of

the antipatharian “black coral” Plumapathes

pennacea (Pallas, 1766) (Cnidaria: Anthozoa),

northwest Gulf of Mexico. Gulf of Mexico

Science 23:127‐132

Bright, TJ, Kraemer GP, Minnery GA, Viada ST

(1984) Hermatypes of the Flower Garden Banks,

northwestern Gulf of Mexico: A comparison to

other western Atlantic reefs. Bulletin of Marine

Science 34:461‐476

Bright TJ, Rezak R (1978) Northwestern Gulf of

Mexico topographic features study. Final Report

to U.S. Department of Interior, Bureau of Land

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION:

TEXAS TO THE FLORIDA STRAITS

  

Sammarco PW, Horn L, Taylor G, Beltz D, Nuttall

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deep tropical coral reef community in the

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Schmahl GP, Hickerson EL, Precht WF (2008)

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communities in the Flower Garden Banks

Region, Northwestern Gulf of Mexico. In: Riegl

BM, Dodge RE (eds) Coral Reefs of the USA,

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Schroeder WW (2007) Seafloor characteristics and

distribution patterns of Lophelia pertusa and

other sessile megafauna at two upper‐slope sites

in the northwestern Gulf of Mexico. U.S. Dept.

of the Interior, Minerals Management Service,

Gulf of Mexico OCS Region, New Orleans, LA.

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Scott‐Denton E, Cryer PF, Gocke JP, Harrelson MR,

Kinsella DL, Pulver JR, Smith RC, Williams JA

(2011) Descriptions of the U.S. Gulf of Mexico

reef fish bottom longline and vertical line

fisheries based on observer data. Marine

Fisheries Reviews 73:1‐26

Silva M, Etnoyer PJ, MacDonald IR (2015) Coral

injuries observed at mesophotic Reefs after the

Deepwater Horizon oil discharge. Deep‐Sea

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Oceanography 129:96‐107

Stiles ML, Harrould‐Kolieb E, Faure P, Ylitalo‐Ward

H, Hirshfield MF (2007) Deep sea trawl fisheries

of the Southeast US and Gulf of Mexico: Rock

shrimp, Royal red shrimp, Calico scallops

Weaver DC, Dennis GD, Sulak KJ (2002) Community

structure and trophic ecology of fishes on the

Pinnacles Reef tract. Department of the Interior,

US Geological Survey, USGS BSR‐2001‐0008

White HK, Hsing P‐Y, Cho W, Shank TM, Cordes

EE, Quattrini AM, Nelson RK, Camilli R,

Demopoulos AWJ, German CR, Brooks JM,

Roberts HH, Shedd W, Reddy CM, Fisher CR

(2012) Impact of the Deepwater Horizon oil spill

on a deep‐water coral community in the Gulf of

Mexico. Proceedings National Academy of

Sciences 109:20303‐20308

White HK, Lyons SL, Harrison SJ, Findley DM, Liu

Y, Kujawinski EB (2014) Long‐term persistence

of dispersants following the Deepwater Horizon

oil spill. Environmental Science and Technology

Letters 1:295‐299

Wicksten MK, Nuttall MF, Hickerson EL (2014)

Crustaceans from antipatharians on banks of the

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Online Annex 1. Deep-sea Coral Taxa in the U.S. Gulf of Mexico Region: Depth and 

Geographic Distribution 

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Appendix 

Map 1. Locations of black corals (Order Antipatharia) recorded in the National Deep‐Sea Coral and Sponge Database (as 

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION: TEXAS TO TH

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Map 2. Locations of stony corals (Order Scleractinia) recorded in the National Deep‐Sea Coral and Sponge Database (as 

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Map 3. Locations of true soft corals (Order Alcyonacea) recorded in the National Deep‐Sea Coral and Sponge Database (

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Map 4. Locations of gorgonian corals (Order Alcyonacea, in part – formerly Order Gorgonacea) recorded in the Nationa

Sponge Database (as of November 2017).

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Map 5. Locations of sea pens (Order Pennatulacea) recorded in the National Deep‐Sea Coral and Sponge Database (as of 

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION: TEXAS TO TH

  

Map 6. Locations of lace corals (Order Anthoathecata, Family Stylasteridae) recorded in the National Deep‐Sea Coral an

(as of November2017).

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STATE OF DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS IN THE GULF OF MEXICO REGION: TEXAS TO TH

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Map 7. Locations of sponges (Phylum Porifera) recorded in the National Deep‐Sea Coral and Sponge Database (as of No

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POPULATION CONNECTIVITY OF DEEP-SEA CORALS

• • •

POPULATION 

CONNECTIVITY OF 

DEEP-SEA CORALS

I. Introduction

Identifying the scale of dispersal among habitats 

has been a challenge in marine ecology for decades 

(Grantham et al. 2003, Kinlan and Gaines 2003, Hixon 

2011). Unlike terrestrial habitats in which barriers to 

dispersal may be obvious (e.g. mountain ranges, rivers), 

few absolute barriers to dispersal are recognizable in the 

sea. Additionally, most marine species have complex life 

cycles in which juveniles are more mobile than adults. As 

such, the dynamics of populations may involve processes 

in distant habitats that are coupled by a transport 

mechanism. Studies of population connectivity try to 

quantify the transport, or dispersal of individuals, among 

geographically separated populations. For benthic marine 

species, such as corals and demersal fishes, colonization 

of new populations occurs primarily by dispersal of 

larvae (Figure 1; Shank 2010). Successful dispersal and 

recruitment, followed by maturation and reproduction of 

these new migrants ensures individuals contribute to the 

gene pool (Hedgecock 2007). Thus, successful dispersal 

links and cohesively maintains spatially separated subpopulations. At shorter time scales (tens to hundreds 

of years), connectivity regulates community structure 

by influencing the genetic composition, diversity and 

demographic stability of the population, whereas at longer 

time scales (thousands of years), geographic distributions 

are affected (McClain and Hardy 2010). Alternatively, 

populations may become extinct or speciation may occur 

if connectivity ceases (Cowen et al. 2007). Therefore, the 

genetic exchange of individuals between populations is 

fundamental to the short-term resilience and long-term 

maintenance of the species. However, for the vast majority 

Cheryl L. Morrison1*, 

Amy R. Baco2, 

Martha S. Nizinski3, 

D. Katharine

Coykendall1,

Amanda W.J.

Demopoulos4,

Walter Cho5, and 

Tim M. Shank6

• • •

1 National Fish Health 

Research Laboratory, U.S. 

Geological Survey, Leetown 

Science Center, WV

* Corresponding Author:

CMorrison@usgs.gov

2 Department of Earth, Ocean, 

and Atmospheric Science, 

Florida State University, 

Tallahassee, FL 

3 NOAA National Systematics 

Laboratory, Smithsonian 

Institution, Washington DC 

4 Wetland and Aquatic 

Research Center, U.S. 

Geological Survey, 

Gainesville, FL

5 Department of Biology, Point 

Loma Nazarene University, 

San Diego, CA

6 Biology Department, 

Woods Hole Oceanographic 

Institution, Woods Hole, MA
of marine species, population connectivity 

remains poorly understood.

To effectively manage marine species, 

knowledge of the spatial scale at which 

populations are connected is beneficial 

(Palumbi 2003, Fogarty and Botsford 2007). 

Ecosystems within the deep sea often occur 

over large geographic scales, yet are spatially 

fragmented across the entire range with 

stretches of unsuitable habitat separating 

prime habitat patches (e.g. cold-water coral 

reefs, methane seeps, hydrothermal vents). 

Habitat fragmentation may escalate with 

increasing resource exploitation (e.g. Hilário et 

al. 2015). The ecological patterns observed in 

many deep-sea ecosystems can be explained 

by metapopulation dynamics as conceived 

by Levins (1969). Under this scenario, species 

occur within patchy habitats that undergo 

local extinction and recolonization and those 

patches are connected via dispersal and 

migration. The persistence, or viability, of the 

metapopulation increases as the availability 

of suitable habitat patches increases, local 

extinction decreases, and migration/dispersal 

between patches increases. In local deepsea communities, the regional distributions 

of each species may be composed of 

metapopulations that are influenced by a 

balance among global-scale, landscape-scale 

and small scale dynamics (Levins et al. 2001).

Figure 1. Diagrammatic representation of potential physical (hydrodynamics, depth) and biological 

(reproduction, larval behavior, settlement) forces that may influence dispersal, colonization, and genetic 

connectivity among deep-sea coral populations. (reprinted from Shank 2010, Oceanography; 1).

380

POPULATION CONNECTIVITY OF DEEP-SEA CORALS

• • •
381

POPULATION CONNECTIVITY OF DEEP-SEA CORALS

• • •

Spatial management tools, such as networks 

of marine protected areas or reserves have 

the potential to protect the entire ecosystem, 

including nursery grounds, adult habitats, 

food sources, etc. Success of these protective 

measures relies at least partially upon 

estimates of dispersal rates across reserve 

boundaries (Botsford et al. 2003, Hilborn et al. 

2004). For example, a population may be more 

vulnerable to human activities if it is not likely 

to receive recruits from other populations 

(e.g., self-sustaining populations, where only 

larvae produced within the population settle 

in the population). Alternatively, if larvae 

produced within a given population disperse 

to other populations, that population may 

be an important larval “source” that could 

rescue depleted populations (Pulliam 1988). 

Identifying source sub-populations is vitally 

important to management and conservation 

plans. Estimates of connectivity provide the 

relative scale at which a protected area may 

function. Such information may be utilized 

to optimize the placement of protected areas 

and to estimate the potential impacts to 

recruitment outside protected area boundaries 

(Palumbi 2001, Gaines et al. 2010).

Biological and physical processes influence 

larval dispersal (Cowen et al. 2007, Paternello 

et al. 2007, Galarza et al. 2009, Cowen and 

Sponaugle 2009, Sivasundar and Palumbi 

2010, Mokhtar-Jamaï et al. 2011, Woodson et 

al. 2012). While biological attributes, such as 

timing of reproduction, larval behavior (e.g., 

swimming, vertical migrations), buoyancy, 

and physiology (e.g., feeding), affect larval 

survivorship and dispersal distances, physical 

processes, such as prevailing currents, eddies, 

recirculating flows, bottom topography and 

upwelling, can enhance or constrain larval 

movement (Figure 1; Shank 2010). Factors such 

as distance from a spawning site, advection 

and diffusion, and high mortality rates, all 

influence the number of larvae present in the 

water column (Cowen and Sponaugle 2009, 

Rosser 2015). The combined influences of 

these biological and physical parameters may 

either promote dispersal of larvae over great 

distances (i.e. create dispersal corridors), or 

constrain population connectivity by isolating 

some populations from others (i.e. create 

barriers to dispersal).

Due to small larval sizes and the immense 

volume of the oceans into which larvae 

disperse, measurements of dispersal distances 

are difficult and tracking of spawning events 

can be quite challenging (Gawarkiewicz et 

al. 2007). Life history traits, such as pelagic 

larval duration, have been used as a proxy for 

dispersal distances. Pelagic larval duration 

correlated well with estimates of dispersal 

for some species (Bohonak 1999, Shanks et al. 

2003). However, there are numerous examples 

where no relationship between pelagic larval 

duration and dispersal distance was observed 

(e.g. Severance and Karl 2006, Miller and Ayre 

2008, Weersing and Toonen 2009, Galarza et 

al. 2009). Although the validity of correlation 

between pelagic larval duration and dispersal 

potential remains equivocal, it may at least 

set an upper bound on dispersal distance 

(Selkoe and Toonen 2011). For the cold-seep 

dwelling mussel “Bathymodiolus” childressi and 

an associated gastropod, Bathynerita naticoidea, 

larvae were detected in surface currents, 

suggesting larvae migrate hundreds of meters 

above the sea floor, allowing greater dispersal 

in faster surface currents (Arellano et al. 2014). 

Unfortunately, for the majority of deep-sea 

coral species, knowledge of larval duration 

and behavior such as swimming ability is 

completely absent. In a review of data on 

pelagic larval durations in deep-sea taxa, a 

total of 21 species have been characterized. 

Of these, only three were cnidarian species 

(Hilário et al. 2015). In fact, knowledge of basic 
382

types of reproduction in deep-sea cnidarians is 

scarce. For example, of the 615 known deepsea scleractinian coral species (Cairns 2007), 

reports on various aspects of reproduction 

have been ascertained for only 15 species 

(Waller 2005, Mercier et al. 2011). The 

structure-forming scleractinian corals Lophelia 

pertusa and Oculina varicosa are seasonal 

broadcast spawning species that release 

gametes into the water column (Brooke and 

Young 2003, Waller and Tyler 2005, Brooke 

and Järnegren 2013). In contrast, hydrocorals 

in the family Stylasteridae, often an important 

structural component of deep-sea coral 

gardens (Stone 2006, Lindner et al. 2008), have 

a reproductive strategy that involves brooded 

larvae with a short planktonic duration, which 

may limit larval dispersal potential (Brooke 

and Stone 2007). Similarly, Corallium rubrum, 

a precious coral with a wide bathymetric 

distribution, also broods larvae and releases 

well-developed, competent larvae (Abbiati 

et al. 2010). In accordance with brooded 

larvae, effective dispersal appears restricted 

(Ledoux et al. 2010a, Constantini and Abbiati 

2015), and shallow water populations are 

genetically structured at a scale of tens of 

meters (Ledoux et al. 2010b, Abbiati et al. 2010, 

Aurelle et al. 2011, Aurelle and Ledoux 2013). 

Many octocorals broadcast gametes into the 

water column annually, relying on external 

fertilization and larvae with an extended 

planktonic development phase (Waller and 

Baco 2007, Kahng et al. 2011, Mercier and 

Hamel 2011, Watling et al. 2011, Waller et 

al. 2014, Nonaka et al., in press, Feehan and 

Waller 2015), so greater dispersal potential 

may be expected. Identification of newlysettled individuals is also extremely difficult, 

and in situ measures of recruitment in deep 

sea corals are limited (Grigg 1988, Thresher et 

al. 2011, Lacharite and Metaxas 2013). 

Traditional methods such as stock assessments 

or analysis of morphological differences 

between populations have been used to 

define conservation management units and 

potential larval sources. However, potential

for connectivity between geographically 

separated populations that do not appear 

to differ morphologically does not mean 

active exchange of larvae is occurring. In 

fact, populations may have been isolated 

for thousands of years or more yet remain 

morphologically similar, resulting in 

genetically differentiated sibling species (see 

Knowlton 1993). Thus, sampling organisms 

from different sites and indirectly measuring 

connectivity using genetic techniques may 

be the only practical way to understand past 

and present connections (Palumbi 2003, Shank 

2010). 

II. Genetic Methods

Use of molecular techniques has changed 

our view of population structuring 

and management. Molecular data have 

demonstrated that a single management 

unit with a homogeneous, wide-ranging 

distribution is not necessarily the best 

management/conservation model (Ayre and 

Hughes 2000, Swearer et al. 2002, Jones et 

al. 2005, Almany et al. 2007). For example, 

the scale over which populations of shallowwater scleractinian corals are differentiated 

can range from 25 to 7,500 km (Baums 2008), 

and may involve differentiation with depth 

(Serrano et al. 2014). A heterogeneous spatial 

mosaic of multiple units with varying amounts 

of genetic structure may better describe what 

is found in nature. Researchers must properly 

sample geographic populations, consult 

taxonomists, use appropriate genetic markers 

(e.g., mitochondrial or nuclear DNA), and 

consult theoretical models to ascertain the 

POPULATION CONNECTIVITY OF DEEP-SEA CORALS

• • •
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POPULATION CONNECTIVITY OF DEEP-SEA CORALS

• • •

genetic structure of a species’ populations 

(Pante et al. 2015a).

Over broad geographic scales (e.g., entire 

ocean basins), mitochondrial DNA markers 

are often employed to assess connectivity. For 

most animals, mitochondrial DNA is inherited 

maternally. Thus, only one copy of each gene 

exists in an individual. Mitochondrial DNA 

markers provide some of the most well-known 

examples of geographic breaks in population 

connectivity among shallow water marine 

species such as oysters, horseshoe crabs, and 

black sea bass along the east coast of Florida 

(Avise 1992). Mitochondrial DNA markers 

have also been useful in identifying closely 

related (sibling) or morphologically similar 

(cryptic) species, both of which commonly 

occur in the deep sea (Rogers 2002). However, 

mitochondrial DNA evolves at a slow rate in 

corals (50-100 times slower than most animals) 

and generally does not provide species-level 

resolution (France and Hoover 2002, Shearer 

et al. 2002, Hellberg 2007, McFadden et al. 

2011, Baco and Cairns 2012). This slow rate of 

mitochondrial DNA sequence evolution has 

impeded studies of connectivity in corals (e.g., 

van Oppen and Gates 2006, Eytan et al. 2009), 

where more variable nuclear DNA markers 

are necessary to assess connectivity.

Realized average dispersal of an organism 

over its entire life cycle (planktonic larval 

stage, settlement, maturation) can be 

estimated from nuclear markers such as 

microsatellites (Neigel 2002). Microsatellite 

markers refer to regions of DNA that include 

adjacent repeats of several nucleotide bases. 

Microsatellites have a higher mutation rate 

than mitochondrial DNA, so often reflect 

connectivity on more recent time scales. 

However, unlike mitochondrial DNA, 

nuclear markers like microsatellites are 

inherited from both parents, so individuals 

have two copies of each nuclear gene, called 

alleles, which are the building blocks that 

populations utilize to adapt and evolve in 

changing environments. These alleles may 

differ in length due to differences in the 

number of times the nucleotide bases are 

repeated. When the alleles are identical, they 

are termed homozygous, different alleles are 

heterozygous. Often, these population-level 

molecular markers are species-specific and 

highly variable. Utilization of rapidly-evolving 

genetic markers with many alleles allows one 

to decipher the unique genetic makeup of each 

individual. Although the initial cost of analysis 

is higher, results with resolution to the level 

of individuals and parentage is often worth 

the expense. Markers with high specificity 

and variability are utilized to assess family 

relationships and clonality in organisms, 

two attributes that complicate the genetic 

signatures of cold-water coral populations.

One of the most important applications 

of population genetic data is to determine 

the level of genetic structuring within and 

between populations. Genetic structure within 

a species can range from very weak, where 

exchange of individuals is common (i.e., high 

connectivity), to strong genetic structuring, 

where exchange of individuals is more 

rare (i.e., low connectivity, moving toward 

isolation of populations; reviewed by Palumbi 

2003, Hedgecock 2007, Cowen and Sponaugle 

2009). Genetic estimates of connectivity result 

from counting differences in DNA sequences 

or examining the variance in allele frequencies 

between populations that accumulate 

when connectivity is low or nonexistent 

(see below). When genetic structuring is 

detected, examination of the physical or 

biological parameters that may be correlated 

to genetic differentiation may provide clues 

to the processes that act to determine spatial 

connections among populations.
384

Population geneticists also attempt to decipher 

patterns of genetic connectivity indirectly 

through the application of theoretical 

models of population structure (Hedgecock 

2007). The stepping-stone model assumes 

neighboring populations are more likely to 

exchange migrants amongst themselves than 

withpopulations further away (Kimura and 

Weiss 1964). Thus, genetic distance should 

increase with geographic distance, populations 

in closer proximity will be genetically more 

similar to each other than to populations 

farther away. This pattern of connectivity, 

referred to as isolation-by-distance (Wright 

1943), creates a highly structured, genetically 

complex system (Rousset 1997). For example, 

an isolation-by-distance pattern was confirmed 

for Porites lobata populations inhabiting 

shallow reefs in the Hawaiian Archipelago 

(Polato et al. 2010). If an isolation-by-distance 

pattern of connectivity is detected, estimation 

of dispersal distances may be biologically 

meaningful (Rousset 1997, Palumbi 2003).

An indirect measure of the persistence of a 

metapopulation is genetic diversity. Genetic 

diversity covaries with the number of 

patches (sources) that can supply migrants 

(Maruyama and Kimura 1980, Vrijenhoek 

2010). Therefore, estimating genetic diversity 

within metapopulations is a proxy for the 

vulnerability of a species to extinction.

III. Connectivity in the Deep

Sea

The majority of studies of marine connectivity 

focus on shallow-water environments, 

particularly tropical reef fishes (Hixon 2011). 

Although shallow marine habitats were once 

considered open systems with ample exchange 

of larvae over large distances, it is now 

accepted that local recruitment and small-scale 

population structure are common despite the 

lack of obvious physical barriers (Cowan and 

Sponagule 2009). Genetic data have suggested 

similar results for shallow-water corals, in 

that most recruitment is local, yet occasional 

long-distance dispersal can occur across tens 

to hundreds of kilometers (Ayre and Hughes 

2000, Miller and Ayre 2008, Gorospe and Karl 

2013).

Although less is known about patterns of 

connectivity in the deep sea, recent studies 

have suggested common themes. First, similar 

to shallow populations along coastlines, 

including many coral species along the Great 

Barrier Reef (Ayre and Hughes 2004), the 

stepping stone model may be appropriate for 

many deep-sea populations, particularly those 

arranged linearly along continental margins 

(LeGoff-Vitry et al. 2004, Smith et al. 2004), 

mid-oceanic ridge axes (Coykendall et al. 2011, 

reviewed by Vrijenhoek 2010) or linear arrays 

of seamounts (Samadi et al. 2006).

In contrast, stretches of open ocean that 

interrupt a linear array of reefs (Ayre 

and Hughes 2004) or vent populations 

(Vrijenhoek 2010) may create an effective 

barrier to dispersal, and connectivity may 

decrease abruptly, creating regionally 

isolated populations. Regional differentiation 

of deep-sea fauna that inhabit continental 

slope habitats may not be limited to benthic 

organisms such as corals. Even mobile species, 

such as fishes (e.g. Helicolenus dactylopterus

(Aboim et al. 2005), squid (Shaw et al. 1999), 

scavenging isopods (France and Kocher 

1996), and red crabs (Weinberg et al. 2003), 

show regional structuring (see Rogers 2002). 

Suggested mechanisms that may prevent 

panmixia of continental slope species 

include physical barriers (continents and 

large expanses of deep water, deep water 

sills (Cowart et al. 2013), or among back-arc 

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POPULATION CONNECTIVITY OF DEEP-SEA CORALS

• • •

basins (Thaler et al. 2014)), structuring of 

the water column (density layers), rugged 

topography (canyons and seamounts) and 

oxygen minimum zones (Rogers 2002). In 

addition, species-specific life history strategies, 

especially related to spawning, may also act 

to limit exchange of individuals between 

populations (Rogers 2002). 

Shared patterns of species diversity and 

genetic structuring across unrelated but 

co-distributed species may indicate that 

oceanographic features affect connectivity of 

many species in similar ways (Cunningham 

and Collins 1998). For example, the deepsea scleractinian coral species Desmophyllum 

dianthus, as well as two antipatharian species, 

Antipathes robillardi and Stichopathes variabilis, 

exhibited genetic subdivision across large 

ocean expanses in the southern Pacific Ocean 

(Miller et al. 2010). Concordance among 

regional connectivity patterns of these codistributed species indicates that physical 

forces (e.g., prevailing currents, eddies, 

upwelling) may restrict larval dispersal among 

regions. However, no genetic subdivision 

was detected for two other coral species 

(Solenosmilia variablis, Madrepora oculata) using 

the same mitochondrial and nuclear gene 

regions (Miller et al. 2010). Conversely, using 

microsatellites instead of mitochondrial DNA, 

Becheler et al. (in review) identified distinct 

genetic populations of Madrepora oculata

among canyons in the Bay of Biscay (Eastern 

North Atlantic Ocean), yet co-occurring 

Lophelia pertusa were panmictic. Clearly, no 

generalized connectivity pattern applies to all 

coral species.

A factor that may uniquely influence 

connectivity among deep-sea organisms is 

depth. Pronounced physiological gradients 

occur as depth increases, which could result 

in locally adapted populations and enhanced 

genetic differentiation (Rogers 2002, Zardus 

et al. 2006). This concept, known as the depthdifferentiation hypothesis (Rex and Etter 

2010), suggests that the divergent selection 

across environmental gradients may cause 

population differentiation, leading to new 

and/or cryptic deep-sea species. Depthrelated divergence has been identified in 

molluscs (e.g. Chase et al. 1998, Etter et al. 

1999, Goffredi et al. 2003, Etter et al. 2005, 

Zardus et al. 2006, Jennings et al. 2013), 

polychaetes (Lundsten et al. 2010, Schüller 

2011, Cowart et al. 2014),amphipods (France 

and Kocher 1996, stylasterid corals (Lindner 

et al. 2008), primnoid octocorals (Baco and 

Cairns 2012), and the red coral Corallium 

rubrum (Constantini et al. 2011). In other 

deep-sea corals, limited vertical larval 

dispersal has been suggested the solitary coral 

Desmophyllum dianthus, a scleractinian coral 

with a cosmopolitan distribution. Genetic 

differentiation with depth in D. dianthus was 

consistent with the stratification of deep 

water masses that entrain larvae and do not 

allow mixing among depth strata (Miller 

et al. 2011). In a study that included three 

species of octocorals in the Gulf of Mexico, 

an ecological niche model revealed little 

overlap among Callogorgia species occupying 

different depths (Quattrini et al. 2013). The 

depth differentiation hypothesis was tested 

of these species, Callogorgia delta, using 

microsatellite data, and isolation by depth was 

confirmed (Quattrini et al. 2015). Rex et al. 

(2005), using deep-sea molluscan fauna with 

bathyal and abyssal distributions as examples, 

postulate that the abyssal environment may 

create population sinks due to a decrease in 

organic carbon influx from more productive 

coastal systems. These isolated populations 

rely on bathyal populations for immigrants. 

Conversely, nine genetic breaks were 

detected in the cosmopolitan deep-sea 
REFERENCE SPECIES REGION MARKERS CONNECTIVITY 

Eytan et al. 2009 Oculina varicosa WNAtlantic 4 nuclear DNA genes Depth structured 

Thoma et al. 2009 6 octocorals, 2 antipatharians WNAtlantic, NE Seamounts mtDNA No structuring, high connectivity 

Cho and Shank 

2010

Asteroschema clavigerum

Ophiocreas oedipus

Ophioplinthaca abyssalis

Ophioplinthaca chelys

WNAtlantic,

NE Seamounts 2 mtDNA genes Regional seamount structure, depth 

Morrison et al. 2011 Lophelia pertusa North Atlantic 9 microsatellites

Regional structure, 

moderate connectivit

within regions

Baco and Shank 

2005

Baco et al. 2006

Corallium lauuense Central Pacific 3-6 microsatellites High connectivity, structure between site

and depths 

Baco and Cairns 

2012 Several Narella spp North Pacific 6 mtDNA markers

Narrow geographic an

depth ranges for 

haplotypes

Herrera et al. 2012 Paragorgia arborea

North Pacific, 

South Pacific, 

North Atlantic

7 mtDNA genes Regional structure 

Quattrini et al. 2015 Callogorgia delta Gulf of Mexico 10 microsatellites Depth structured 

Table 1. Studies examining connectivity among deep-sea corals and/or deep coral communities in U.S. waters. 

386

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POPULATION CONNECTIVITY OF DEEP-SEA CORALS

• • •

amphipod Eurythenes gryllus. In this instance, 

higher diversity was observed at abyssal 

rather than bathyl depths, thus reflecting a 

pattern contrary to the depth-differentiation 

hypothesis (Havermans et al. 2013). Clearly, 

environmental factors associated with depth 

may create important abiotic gradients that 

may influence population structuring in the 

deep sea (Quattrini et al. 2015).

IV. Connectivity Among

Deep-Sea Corals in U.S.

Waters

IV.1. North Atlantic Ocean

Genetic methods were used to assess 

connectivity patterns in several North Atlantic 

deep-sea coral species (Table 1). In the first 

of these studies, connectivity among shallow 

and deep populations of the structureforming coral Oculina varicosa was examined 

using three nuclear DNA sequence markers 

(Eytan et al. 2009). Oculina varicosa occurs 

most commonly at shallow depths (< 30 m). 

However, azooxanthellate colonies grow at 70 

to 100 m depth along the Oculina Banks off the 

Florida east coast (Reed 2002). Despite Federal 

protection, the Oculina Banks have been 

negatively impacted by illegal trawling and 

dredging (Reed et al. 2007). The Oculina Banks 

population was found to be distinct from 

shallow populations; therefore, depth was 

considered an important factor structuring the 

pattern of connectivity in O. varicosa (Eytan et 

al. 2009). 

Lophelia pertusa, a structure-forming coral 

species commonly found on the continental 

slope off the southeastern U.S. coast (Ross 

and Nizinski 2007) and Gulf of Mexico 

(Brooke and Schroeder 2007), has a nearly 

cosmopolitan distribution, suggesting 

substantial dispersive ability. Lophelia pertusa 

is a broadcast spawner that was originally 

thought to produce non-feeding larvae (Waller 

and Tyler 2005). Yet a recent study suggests 

larvae may feed and actively swim upwards 

during the first weeks after fertilization, a 

behavior that may allow larvae to rise above 

the benthic boundary layer and promote 

advection in stronger currents (Larsson et 

al. 2014). Swimming behavior as well as an 

estimated three to five week larval duration 

imply high dispersal potential. However, 

the spatial scales that L. pertusa larvae 

travel remains unknown, making accurate 

predictions of realized larval dispersal 

challenging. Nine microsatellites were used 

to examine patterns of genetic connectivity 

across a large portion of the geographic range 

of L. pertusa in the North Atlantic Ocean 

(Morrison et al. 2011). Four distinct genetic 

groupings corresponding to ocean regions 

were identified: Gulf of Mexico, coastal 

southeastern U.S., New England Seamounts, 

and eastern North Atlantic Ocean (Figure 

2). It is known that reproductive timing is 

offset between the eastern North Atlantic 

(Waller and Tyler 2005) and Gulf of Mexico 

(Brooke et al. 2007) L. pertusa populations. 

Whether or not regionally isolated L. pertusa 

populations represent cryptic species will 

require further morphological examination 

and biological evidence. Interestingly, 

similarities exist between zoogeographic 

patterns of deep-sea scleractinian corals 

(Cairns and Chapman 2001) and regional L. 

pertusa genetic differentiation (Morrison et 

al. 2011), suggesting that similar mechanisms 

(see above) may constrain coral larvae within 

regions. 

Although the populations of L. pertusa at 

the New England seamounts occur in a 

geographic location that could be a corridor, 

or stepping stone, for connectivity across the 

Atlantic Ocean, the seamount populations 
388

were the most genetically differentiated 

based upon microsatellites, suggesting 

little connectivity (Morrison et al. 2011). 

The seamount populations appeared more 

genetically similar to populations from the 

eastern rather than western North Atlantic 

Ocean, despite occurring in closer proximity to 

the latter. These seamount populations occur 

deeper (1418-1679 m) than other populations 

sampled (140-740 m) and therefore differing 

water masses and circulation patterns may act 

as a barrier to gene flow.

For North Atlantic Ocean L. pertusa 

populations, a positive correlation between 

genetic and geographic distance was detected 

at broader scales (thousands of km), but not 

at smaller scales (Morrison et al. 2011). Since 

regional breaks in connectivity were also 

detected, deductions regarding dispersal 

distances may be compromised due to the 

likely violation of the stepping-stone model 

assumption of on-going dispersal at these 

large scales (Slatkin 1993, Garnier et al. 

2004). However, results suggest moderate 

connectivity within each of the regions. A 

Figure 2. Probability of assignment of Lophelia pertusa individuals, represented by colored vertical 

bars, to four groups based upon multi-locus genotypes using the program STRUCTURE. Populations are 

arranged from west (left) to east (right). The top bar (round 1) represents an initial run including all L. 

pertusa samples. The lower bar represents additional structuring between New England Seamounts and 

Eastern North Atlantic populations detected during a subsequent run of STRUCTURE. (Reprinted from 

Morrison et al. 2011, Conservation Genetics; 37). 

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POPULATION CONNECTIVITY OF DEEP-SEA CORALS

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previous study of L. pertusa in the eastern 

North Atlantic Ocean also reported moderate 

connectivity (genetic cohesion) among 

European continental margin populations 

and no correlation between genetic and 

geographic distance (Le Goff-Vitry et al. 

2004). Levels of both inbreeding and asexual 

reproduction (clones) varied among European 

margin L. pertusa populations, suggesting 

that most dispersal is spatially restricted, yet 

occasional long-distance dispersal is adequate 

to maintain genetic cohesion (Le Goff-Vitry et 

al. 2004). While both studies detected genetic 

structuring (i.e., decreased connectivity) 

between continental slope and Norwegian 

fjord L. pertusa populations in the eastern 

North Atlantic, results from Morrison et al. 

(2011) suggest the magnitude of differentiation 

is less than that observed across the North 

Atlantic Ocean. 

Lophelia pertusa populations surveyed in 

both the eastern and western North Atlantic 

Ocean using microsatellites have shown 

heterozygote deficiencies (Le Goff-Vitry 

et al. 2004, Morrison et al. 2011, Dahl et al. 

2012). Similarly, heterozygote deficits have 

been detected in the majority of population 

genetic studies of corals (Ayre and Hughes 

2000, van Oppen and Gates 2006, Selkoe 

and Toonen 2006), including cold-water 

corals (Baco and Shank 2005, Baco et al. 

2006). Occurrence of heterozygote deficits 

in nature is often explained as a violation 

of the concept of the ‘ideal population’ 

assumed by the Hardy-Weinberg principle. 

For instance, a subset of the population 

could contribute the most genetic material 

to the next generation (non-random mating). 

Studies have shown a propensity for clonal 

reproduction via fragmentation in most L. 

pertusa populations surveyed (Le Goff-Vitry et 

al. 2004, Morrison et al. 2011, Dahl et al. 2012). 

Clonal reproduction can cause heterozygote 

deficits due to deviation from random mating 

(Dahl et al. 2012, van Oppen et al. 2008). In 

fact, asexual reproduction via fragmentation 

may play a key role in the establishment and 

maintenance of deep L. pertusa reefs (Dahl et 

al. 2012). For example, few genetic individuals 

become dominant at a reef site and skew 

opportunities for mating. Additionally, 

the lifespan of a genetic individual may be 

quite long (thousands of years; Dahl et al. 

2012), therefore, this life history trait may be 

important to both estimates of connectivity 

and to the evolution of the species. 

Seamounts are hypothesized to be locations 

of isolation in the deep-sea, resulting in 

high levels of speciation and endemism 

(e.g. Hubbs 1959, Rowden et al. 2010, Pante 

et al. 2015b). Three studies (Thoma et al. 

2009, Cho and Shank 2010, Herrera et al. 

2012) have addressed connectivity along the 

New England and Corner Rise Seamounts. 

Thoma et al. (2009) investigated isolation 

among individual seamounts by testing 

the hypothesis that corals occurring at 

isolated seamounts would possess unique 

mitochondrial DNA haplotypes. Variability 

in the mitochondrial mismatch repair gene 

homolog (mutS) was assessed for five octocoral 

genera. Nine markers for antipatharians as 

well as an intergenic spacer for bamboo corals 

were also incorporated into the data set. 

The Thoma et al. (2009) study found four 

haplotypes unique to the seamounts, as well 

as two haplotypes from the adjacent margin 

that were widespread, consistent with the 

possibility of seamounts being hotspots for 

divergence of species. A caveat of this work 

is that the unique haplotypes discovered 

at the seamounts were not necessarily the 

result of endemism, but could be due to small 
390

sample sizes. This work, however, is based 

on the assumption that mutS haplotypes were 

species-specific. Subsequent work has shown 

this is not the case. Rather, some species of 

octocorals have multiple haplotypes for mutS, 

which can lead to underestimating species 

ranges, while other mutS haplotypes are 

shared between multiple species, resulting 

in overestimating species ranges (McFadden 

et al. 2010, McFadden et al. 2011, Baco and 

Cairns 2012, Pante et al. 2015b). These same 

studies show that only about 50% of species 

could be resolved using the mutS marker. 

Thus the single-marker approach of Thoma 

et al. (2009) likely does not provide sufficient 

power to resolve the connectedness of the New 

England and Corner Rise seamounts at the 

species level.

In contrast, data from seven mitochondrial 

gene regions and nuclear genetic variants 

of the deep-sea bubblegum coral Paragorgia 

arborea revealed basin-wide and global 

patterns of genetic variation correlated 

with historical migrations and connectivity 

from the Western Pacific into the Atlantic 

(Herrera et al. 2012). Prior to this study, 

diversity throughout the entire known 

distribution of a cold-water coral species had 

not been evalutated. Herrera et al. (2012) 

utilized mitochondrial and nuclear genetic 

variants in a phylogeographic context to 

examine the compatibility of P. arborea with 

the genealogical-phylospecies concept by 

examining specimens collected over its 

known distribution. The multi-marker use 

demonstrated that the global morphospecies 

P. arborea can be defined as a genealogicalphylospecies hosting differing levels of

connectivity around the worlds’ oceans.

Global genetic variation among populations

of this species revealed significant basin

scale differences. These phylogeographic

data suggest a scenario in which P. arborea

originated in the North Pacific, possibly 

in the Western North Pacific, followed 

by colonization of the South Pacific and 

spreading eastward around the Southern 

Hemisphere in a stepping stone fashion 

(possibly via the Antarctic Circumpolar 

Current). The colonization of the North 

Atlantic seems to have occurred through a 

more recent dispersal event from the South 

Pacific, via the Central American Seaway, 

or from the South Atlantic. Notable is that 

despite the finding of significant geographic 

variation over the geographic range, no 

significant correlation with depth was 

observed.

A study by Cho and Shank (2010) 

demonstrated the impact host-specific 

symbiotic relationships may have on the 

genetic connectivity of fauna associated with 

cold-water corals. Deep-sea corals provide 

habitat for many species that have developed 

symbiotic relationships with varying degrees 

of specificity to their host corals (Shank 2010, 

Buhl-Mortensen et al. 2010). The co-evolution 

of host-associate relationships may affect the 

genetic connectivity of these coral associates. 

Cho and Shank (2010) studied the patterns 

of dispersal and genetic connectivity of four 

brittle star species (Asteroschema clavigerum, 

Ophiocreas oedipus, Ophioplinthaca abyssalis and 

Ophioplinthaca chelys; Figure 3) that display 

differing levels of associative specificity to 

deep-sea coral hosts inhabiting the New 

England and Corner Rise seamounts.

Analyses of two mitochondrial markers, 

mt16S and mtCOI, revealed species-specific 

genetic differentiation based on geography 

and depth. Asteroschema clavigerum and 

O. oedipus showed significant isolation by

distance, significant genetic differentiation

by depth, and predominantly westward

historical migration. Asteroschema clavigerum

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POPULATION CONNECTIVITY OF DEEP-SEA CORALS

• • •

had significant genetic differentiation by 

geographic region. In contrast, O. abyssalis and 

O. chelys displayed predominantly eastward

historical migration and O. chelys displayed

significant genetic differentiation within

individual seamounts. The patterns of genetic

differentiation shared by A. clavigerum and O.

oedipus may be correlated with species-specific

host specificity and dispersal strategies. Both

of these species have seemingly obligate

symbiotic relationships to their specific coral

hosts and are broadcast spawners, while the

other two species studied, O. abyssalis and O.

chelys, have lower or no fidelity to their coral

hosts and are potentially brooders.

IV.2. North Pacific Ocean

Although scleractinian reefs are not common 

in the North Pacific Ocean, dense deep-sea 

coral assemblages, dominated by octocorals 

and antipatharians, are found throughout 

the region (Baco 2007, Hourigan et al. 2007). 

Despite the high diversity and abundance of 

corals, population genetics analysis of only 

one species has been published. Hemicorallium 

laauense (= Corallium laauense), a commercially 

important octocoral in the family Coralliidae, 

harvested periodically as part of the precious 

coral fishery (reviewed in Grigg 2002), is 

numerically dominant in the Hawaiian 

Archipelago at 350-575m depth (Parrish and 

Baco 2007, Baco 2007). Hemicorallium laauense

appears to spawn with a periodic or quasicontinuous reproductive strategy (Waller and 

Baco 2007).

Figure 3. Images of four brittle star species studied by Cho and Shank (2010). A) Brittle star Asteroschema clavigerum found on the deep-sea coral Paramuricea sp. B) Brittle star Ophiocreas oedipus found 

on the deep-sea coral Metallogorgia melanotrichos. C) Brittle stars Ophioplinthaca abyssalis found 

on the deep-sea coral Candidella imbricate. D) Brittle stars Ophioplinthaca chelys found on the sea 

floor. Image credit: WHOI; DASS 2005 Expedition; IFE; NOAA OE.

A B

C D
392

Because of the relatively linear nature of the 

Hawaiian Archipelago, one would expect a 

classic isolation-by-distance signature in the 

genetic data for most species. However, results 

from an analysis of three microsatellite loci 

in H. laauense, collected from eight sites in the 

Main Hawaiian Islands and southern end of 

the northwestern Hawaiian Islands, indicated 

no isolation-by-distance pattern (Baco and 

Shank 2005). Instead, results indicate that the 

scale of genetic structure on isolated features 

such as oceanic islands and seamounts may be 

more complex than a simple stepping-stone 

model. For example, the deepest site, located 

off Kauai in the middle of the sampling area, 

was quite different genetically compared to 

the majority of sites sampled during the study. 

Also, there was significant variation within 

the continuous Makapu’u bed (on the scale of 

1.6 km) off the Island of Oahu. The fairly high 

heterozygote deficiency found for this species 

based on six microsatellite markers (Baco and 

Shank 2005, Baco et al. 2006) could indicate 

a high degree of inbreeding for this species, 

implying significant isolation for each of the 

precious coral bed locations investigated, with 

only occasional long distance dispersal events. 

These results suggest connectivity within and 

between each feature on or each location along 

a seamount chain should be assessed when 

considering any management actions.

Analyses based upon mitochondrial DNA 

also provide some insight into connectivity 

along North Pacific seamounts. Previous 

studies of seamount octocorals indicated 

widely distributed mitochondrial haplotypes 

and counter the idea that seamounts may be 

isolated (Smith et al. 2004, Thoma et al. 2009). 

However, each of these studies cautions that 

mtDNA markers were not morphologically 

ground-truthed to determine the level of 

genetic variation which corresponds to a 

species. A recent study of the octocoral genus 

Narella (Family Primnoidae) examined the 

variability of six mtDNA regions relative 

to species designations in this genus (Baco 

and Cairns 2012). The results of this study 

indicated that the markers used in previous 

studies could not distinguish individuals 

at the species level (mutS) and two markers 

(indel regions) could not distinguish between 

genera (Smith et al. 2004) based on mtDNA 

haplotypes. Thus, the fact that haplotypes 

were widespread is not sufficient evidence to 

disprove isolation of seamount features. Using 

all six mtDNA markers on specimens collected 

on North Pacific seamounts, the results of the 

Narella study indicate that geographic and 

bathymetric ranges of seamount species may 

be much narrower than previously thought.

V. Future Directions

Whereas the examples discussed above 

provide insights into patterns of connectivity 

for several abundant and important deepsea coral species in U.S. waters, much work 

remains before we can achieve a fundamental 

understanding of the factors that control 

connectivity among coral species and their 

diverse associated fauna. Several specific 

research questions that may guide future 

studies of connectivity among deep-sea corals 

are suggested below.

V.1. How Far do Larvae of DeepSea Corals and Associated

Organisms Disperse?

A basic understanding of distributions 

of deep-sea coral species and associated 

fauna, coupled with a thorough taxonomic 

evaluation, is necessary to address this 

research question (see Pante et al. 2015a). 

Although general ranges for many deep-sea 

coral species are known, the need exists to 

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POPULATION CONNECTIVITY OF DEEP-SEA CORALS

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fine-tune estimates and fill in information 

gaps by visiting unexplored areas. Exploratory 

cruises are necessary to locate populations, 

assess abundances, and collect samples. 

Success in locating and sampling deep-sea 

corals and associated organisms during 

exploratory cruises will depend upon access 

to deep-submergence technologies (see 

below). Additionally, such explorations 

may be guided by predictive biophysical 

modeling, which may help locate where 

previously unknown coral habitats occur 

(e.g. Yearsley and Sigwart 2011). Predictive 

modeling involves methods that produce 

distribution maps from limited sampling 

data, as interpreted through remote sensing 

information such as multibeam bathymetry, 

substrate and geomorphology, rugosity, slope 

and aspect (e.g., Howell et al. 2011) and/or 

aspects of the environmental tolerances of 

corals compared with physical, chemical and 

biological variables at the geographic locations 

being studied (Davies et al. 2008, Guinotte 

et al., this volume)). The effective use of 

biophysical models for deep-sea organisms is 

hampered by knowledge gaps in both physical 

and biological parameters (Hilário et al. 2015

While sampling deep-sea organisms can be 

labor intensive and expensive, obtaining 

sufficient numbers of individuals per 

population for statistical testing has strongly 

impeded progress to date. Need for adequate 

numbers of samples is regarded as essential 

and of critical importance. A balance between 

adequate sampling of local populations and 

sampling broadly throughout the geographic 

range of the species is imperative (Pante et 

al. 2015a). Therefore, funding agencies and 

science parties need to appreciate the value of 

connectivity studies and dedicate dive time 

specifically for sampling spatially structured 

populations. Much can be gained through 

collaborative efforts in large-scale and multispecies connectivity studies if adequate time 

for intensive sampling of individual species is 

included in the sampling plan.

With the advance of new genetic tools and 

the unknown scales of dispersal for almost 

all deep-sea corals and their symbionts, finescale (spatial and temporal) studies are now 

needed. For example, genetic structure was 

found at very small spatial scales (within a 

reef) but disappeared or became negative at 

larger scales in Hawaiian Pocillopora corals 

(Gorospe and Karl 2013). Clonal reproduction 

through fragmentation in scleractinian corals 

may be key to extreme longevity of a genetic 

individual (clone), and may skew chances 

for reproduction to a few individuals (Dahl 

et al. 2012). Therefore, sampling should 

address fine-scale structuring adequately. 

The feasibility of such a sampling scheme has 

been demonstrated (Becheler et al., in review). 

Larger sample sizes and additional markers 

are necessary to obtain the statistical power 

necessary to distinguish small but important 

genetic distinctions (Waples 1998, Hedgecock 

2007). 

V.2. Can Larval Dispersal

Distances be Accurately

Estimated Given Current

Knowledge of Early Life Histories?

Early life history traits, such as timing 

and potential seasonality of reproduction, 

reproductive output and success, planktonic 

duration, and recruitment, all influence 

connectivity (e.g. Underwood and Fairweather 

1989, Rosser 2015). Reproductive periodicity, 

larval type and behaviors are fundamental 

processes that need further investigation 

(Hilário et al. 2015). In the few species that 

have been studied, nearly all modes and 

patterns of reproduction have been observed 
394

(Patarnello et al. 2007, Metaxas and Kelly 2010, 

Mercier et al. 2011). Thus, making predictions 

on dispersal, based upon taxon, depth, or 

location, is extremely difficult.

Assessing patterns of connectivity among 

organisms associated with deep-sea coral 

habitats will expand understanding of the 

functioning of and potential sensitivities 

to disturbance of these ecosystems. By 

comparing patterns of genetic connectivity 

between different species within the same 

habitat, we gain insight into what drives 

migration and dispersal in these ecosystems 

(community genetics). Contrasting patterns of 

population structure observed among species 

within the same habitat indicates different 

life histories are driving resultant dispersal 

patterns (i.e., vagile vs sessile, lecithotrophic 

vs planktotrophic larvae) whereas similar 

patterns of population structure among 

species within the same habitat indicate 

larger forces, such as deep ocean current 

circulation that entrain larvae, are acting on 

organisms similarly. For example, Samadi 

et al. (2006) examined connectivity among 

populations of several squat lobster species 

and two gastropod species along the Norfolk 

ridge seamounts. Results suggested that 

connectivity is restricted only for species with 

limited dispersal ability. Thus, life history 

is an important factor shaping connectivity 

patterns along this seamount chain.

V.3. To What Extent do Patterns

of Connectivity Relate to

Hydrodynamics?

For shallow-water marine taxa, integration 

of genetic, biological, geographic and 

hydrodynamic data into dispersal models 

has provided both estimates of connectivity 

as well as descriptions of physical forces that 

likely shape connectivity patterns (Manel et 

al. 2003, Galindo et al. 2006, Cowen et al. 2006, 

Baums et al. 2005, Baums et al. 2006, Werner 

et al. 2007, Mokhtar-Jamaï et al. 2011, Foster et 

al. 2012). Hydrodynamic patterns surrounding 

deep-sea coral areas are less known (see 

Hilário et al. 2015, for review). However, 

physical data are now being collected in some 

deep-sea regions through the deployment 

of instrumentation that sequentially collects 

environmental data over either short 

(autonomous underwater vehicles) or long 

periods of time (benthic landers; e.g. Davies 

et al. 2010, Yearsley et al. 2011). Integration of 

ocean circulation and larval transport models 

(based upon laboratory tested larval durations 

of seven deep-sea invertebrates) exemplified 

the various patterns of dispersal possible 

(Young et al. 2012). General conclusions from 

this study were that most larvae are retained 

in the same geographic location as adults, 

but when there is net transport of larvae out 

of an area, it is unidirectional in an eastward 

and northward trajectory in Intra-American 

seas (Young et al. 2012). The combination of 

genetic connectivity and hydrodynamic data 

at key time periods (i.e., coral spawning) 

should allow for better precision in predictive 

modeling of both dispersal potential and coral 

presence.

V.4. What Research Tools May be

Useful for Studies of Connectivity

in the Deep Sea?

Continued development of genetic tools 

is necessary to fill large knowledge gaps 

regarding evolutionary processes that shape 

diversity in the deep sea. Through the 

application of genetic techniques, patterns 

and limits of species distributions as well as 

patterns of dispersal, migration, recruitment 

and clonality can be described. Choosing 

the most appropriate genetic tools is projectPOPULATION CONNECTIVITY OF DEEP-SEA CORALS

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• • •

specific, dependent upon the question 

driving the research. For poorly characterized 

taxa, DNA sequence data is necessary to 

identify and refine species boundaries and 

relationships between closely related taxa 

prior to further analyses (Knowlton 2000). 

For example, examination of morphological 

and phylogenic data is necessary prior to 

population genetic analyses for most octocoral 

species (McFadden et al. 2010). Given the 

prevalence of regional and bathymetric 

cryptic species (France and Kocher 1996, 

Etter et al. 1999, Etter et al. 2005, Chase et 

al. 1998, Vrijenhoek 2009, Miller et al. 2010, 

Miller et al. 2011, Puillandre et al. 2011, 

Mantelatto et al. 2014, Pante et al. 2015a), it 

is likely that the number of deep-sea species 

has been underestimated (Rogers 2002). 

Broad-scale sampling and DNA sequencing 

of few samples can address the taxonomy as 

well as provide a first order understanding 

of relationships among geographically 

distant populations. Adequate sampling, 

plus simultaneous molecular and traditional 

systematic investigations may eventually 

produce more accurate species lists and better 

evolutionary descriptions of the evolutionary 

relationships among them (Vrijenhoek 2009, 

Pante et al. 2015a). These kinds of results 

would provide the foundation for future 

population genetic studies. Sampling of 

this nature should be incorporated into 

cruise objectives. Additionally, continued 

database development, promotion of data 

and sample sharing, plus synthesis and 

collaboration, should allow for advancements 

in understanding (McClain and Hardy 2010). 

Inexpensive production of large volumes 

of sequence data for any organism is now 

possible using next-generation sequencing 

technologies (NGS). These new sequencing 

technologies offer exciting prospects 

for marker development in deep-sea 

coral taxa. Researchers using NGS can 

identify microsatellites more quickly and 

inexpensively (e.g. Coykendall and Morrison 

2013, Morrison et al. 2015). Additional types 

of molecular markers, such as single-copy 

nuclear genes (e.g. Concepcion et al. 2008), 

and single-nucleotide polymorphisms 

(SNPs, e.g. Baird et al. 2008), may now be 

developed without prior knowledge of the 

genome (Reitzel et al. 2013). SNPs are often 

numerous (hundreds to thousands) and 

may provide higher genome coverage than 

microsatellites (tens of markers). The utility 

of SNPs for delimiting recalcitrant species of 

octocorals in the genera Chrysogorgia (Pante 

et al. 2015c) and Paragorgia (Herrera and 

Shank 2015) was recently demonstrated. 

Whether or not differentiation is detected 

using neutral genetic markers as discussed 

thus far, populations may be adapted to local 

environmental conditions. By sequencing 

coding regions of the genome, genes that may 

underlie functional variation can be identified 

and surveyed through gene expression 

analyses (e.g. Vera et al. 2008). As such, 

advances in both sequencing technologies 

and statistics for population genomics may 

allow researchers to identify genome-wide 

signatures of adaptation (Baums 2008, Ledoux 

et al. 2015). Collection of environmental data 

that may be correlated with differences in 

gene expression will add value to population 

genomic studies and may help to identify 

the mechanisms that generate and maintain 

diversity. 

For certain deep-sea coral associates, 

chemistry of calcified structures (e.g., fish 

otoliths and molluscan statoliths) can be 

used to assess natal origins of individuals 

and help define geographic scale of dispersal 

and connectivity. These metabolically 

inert structures record the chemistry of the 

environment in which the animal lives. 
396

Natural tags are derived from variation 

in environmental conditions, including 

temperature, salinity, dissolved oxygen, 

and other chemical parameters. If the 

environmental chemistry is distinct, then these 

chemical tags provide a record that chronicles 

spatial separation during spawning, and 

differences between populations (Campana 

et al. 1994). While connectivity in shallowwater environments has been addressed using 

this technique (Thorrold et al. 2007, Eldson 

et al. 2008), application in the deep sea has 

been limited to a few studies focused on 

stock assessments of commercially exploited 

fishes (Hauser and Carvalho 2008). Combined 

approaches using microsatellite markers and 

otolith chemistry can help assess fine scale 

genetic structure in deep-sea fishes, because 

these tools document both the ecological 

and evolutionary timescales appropriate for 

assessing dispersal (Carlsson et al. 2011)

Working in the deep-sea environment is 

challenging and costly, and the advancement 

of connectivity research and exploration 

is intimately tied to advancements in 

deep-submergence technologies. Future 

technological achievements will be those 

that comprehensively expand our ability 

to characterize factors fundamental to 

understanding connectivity, including 

refining distributions of subpopulations and 

habitats, obtaining co-located oceanographic 

data over relevant temporal and spatial 

scales of connectivity, confirming the 

presence of fauna and habitat availability, 

and collecting sufficient numbers of 

population-level samples for taxonomy 

and connectivity studies. Improved AUVs 

offer long-range multi-sensor platforms for 

spatially-expansive survey coverage, precise 

dynamic navigation, fine-scale bathymetric 

mapping, high-resolution photographic 

and chemical laser imaging (e.g., detecting 

species-specific chemicals secreted by corals), 

coupled with coincident oceanographic 

data. The use of remotely-deployed cameras 

(e.g., drop cameras and towed camera sleds) 

allows for rapid ground-truthing of habitats 

and targeting of prime habitat for further 

observation, exploration, and sampling 

using Remotely Operated Vehicles (ROVs) 

and/or submersibles. Development of novel 

sampling gear for ROVs and submersibles 

that allows many discrete samples to be 

collected per dive will increase sampling 

resolution and extent. Lastly, strategicallyplaced long-term monitoring instrumentation, 

including time-lapse cameras, oceanographic 

data samplers, plankton samplers and nonlethal sample collection devices, will help 

resolve long-term oceanographic patterns 

as well as provide insights into species 

behaviors and larval transport. Advancing 

technological capabilities, including 

autonomous oceanographic characterization 

of seafloor habitats, expands our fundamental 

understanding of benthic processes and their 

link to coral community connectivity.

VI. Conclusions

Understanding connectivity and the 

processes that influence connectivity can 

enhance the conservation of deep-sea coral 

diversity through science-based stewardship. 

Connectivity through exchange of larvae 

among populations should be one of the 

factors considered during the planning and 

design phases of deep-sea coral protection 

areas. Combining estimates of genetic 

connectivity with physical oceanographic 

data will lead to a better understanding 

of processes that underlie larval dispersal 

and, therefore, connectivity. Population 

connectivity data among deep-sea coral 

populations is slowly becoming available. 

However, more work is needed and research 

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POPULATION CONNECTIVITY OF DEEP-SEA CORALS

• • •

must continue as scientists and managers 

work toward protecting and conserving these 

ecologically valuable ecosystems.

VII. Acknowledgements

The authors would like to thank Rhian Waller 

(University of Maine), Than Hitt (USGS), Tom 

Hourigan (NOAA), and Peter Etnoyer (NOAA) 

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Squat lobster perched on Lophelia pertusa colonies with a sponge in the background. Courtesy

of NOAA/USGS.

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Thomas F. 

Hourigan1*, 

John Reed2, 

Shirley 

Pomponi2, 

Steve W. Ross3, 

Andrew W. 

David4, and 

Stacey Harter4

  

1 NOAA Deep Sea Coral

Research and Technology

Program, Office of Habitat

Conservation, Silver

Spring, MD  

* Corresponding Author:

Tom.Hourigan@noaa.gov

2Marine Biomedical and

Biotechnology Research

Program, Harbor Branch

Oceanographic Institute,

Florida Atlantic University,

Fort Pierce, FL

3 Center for Marine Science,

University of North

Carolina, Wilmington

4 Sustainable Fisheries

Division, NOAA Southeast

Fisheries Science Center,

Panama City, FL

                   

                 

               

                 

                         

                 

                 

                 

                   

     

STATE OF THE DEEPSEA CORAL AND 

SPONGE ECOSYSTEMS 

OF THE SOUTHEAST 

UNITED STATES 

I. Introduction

The Southeast U.S. region stretches from the Straits of Florida

north to Cape Hatteras, North Carolina, and encompasses the

Southeast U.S. Continental Shelf large marine ecosystem (LME;

Carolinian ecoregion) and associated deeper waters of the Blake

Plateau, as well as a small portion of the Caribbean LME off the

Florida Keys (eastern portion of the Floridian ecoregion). Within

U.S. waters, deep‐sea stony coral reefs reach their greatest

abundance and development in this region (Ross and Nizinski

2007). This warm temperate region is strongly influenced by the

northern‐flowing Gulf Stream.

Along with the Gulf of Mexico, this region has been the focus of

some of the most extensive U.S. deep‐sea coral research, yet many

of the region’s deeper waters remain poorly explored. The South

Atlantic Fishery Management Council (SAFMC) has authority over

most fisheries in Federal waters in this region, which includes the

waters off North Carolina, South Carolina, Georgia, and the

Atlantic coast of Florida, including the Florida Keys. In 1984 the

SAFMC recommended, and NOAA’s National Marine Fisheries

Service (NMFS) established, the Oculina Bank Habitat Area of

Particular Concern (HAPC) —the world’s first marine protected

area specifically designed to protect deep‐sea corals. Over the last

decade, NOAA and the Council have built on these initial steps to

craft a truly comprehensive approach to conserving vulnerable

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

deep‐sea coral habitats by expanding the

Oculina Bank HAPC and establishing five

additional deepwater Coral Habitat Areas of

Particular Concern (C‐HAPCs) on the

southeastern U.S. continental margin covering

59,560 km2 (see http://safmc.net/managed‐

areas/deep‐water‐corals).

In this chapter, we highlight new scientific

information on deepwater coral ecosystems that

has become available since the 2007 deep‐sea

coral status report, and review new

management measures that help protect these

habitats in the region. We also include

information on deepwater sponges, which were

not addressed in the 2007 report.

II. Update on Scientific

Information

II.1. New Research – Overview

Since the research summarized by Ross and

Nizinski (2007), there have been a number of

surveys of deep‐sea coral habitats in the region

(Figure 1; Appendix Table A). Much of this

effort has focused on deep‐sea coral mounds off

eastern Florida, which are dominated by the

coral Lophelia pertusa. There have also been

significant new surveys of shallower shelf‐edge

habitats associated with new snapper‐grouper

marine protected areas (MPAs). These areas

contain rocky hardgrounds with coral

(primarily gorgonian) and sponge assemblages

that were not included in the 2007 report and

are quite different from those that occur in

deeper water.

In 2009, NOAA’s Deep Sea Coral Research and

Technology Program chose the southeast region

for its first 3‐year field research initiative to

locate, study, and provide information for the

conservation of deep‐sea coral and sponge

habitats. The initiative was developed in

consultation with the SAFMC and surveys

targeted information needed for management.

From 2009 – 2011, fieldwork employed a three‐

tiered approach: 1) systematic reconnaissance

mapping using multibeam sonar to provide the

physical context for deep‐sea coral habitats; 2)

targeted high‐resolution mapping; and 3) in situ

ground‐truthing and multidisciplinary research

using submersibles or remotely operated

vehicles (ROVs) to understand the distribution

and ecological role of deep‐sea corals. The

Southeast Deep‐Sea Coral Initiative, in

partnership with numerous research

institutions, supported seven major research

cruises, including two dedicated solely to

multibeam mapping, two dedicated to

remotely‐operated vehicle (ROV) or

submersible surveys, and three combined ROV

and mapping missions (Figure 1; Annex 1).

Over 10,000 km2 of seafloor were mapped.

There were also 22 submersible dives, 36 ROV

dives, numerous midwater and benthic samples

and oceanographic measurements collected.

The initiative also developed predictive habitat

models for deep‐sea corals (see below), which

complemented the field research.

In addition to the Deep Sea Coral Research and

Technology Program, NOAA’s Coral Reef

Conservation Program supported annual

research cruises from 2012‐2015 designed to

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Figure 1. Map showing deep‐water dives (> 50 meters) from Southeast U.S. region in years 2007‐2016.

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Table 1. Number of coral species reported from the Southeast U.S. in 2007 and in the present report. Increases

include new species identifications and range extensions, but principally represent species not included in the

first report. *Not confirmed from U.S. waters.

Taxon Ross and Nizinski 2007 This Report

Class Anthozoa

Black corals (Order Antipatharia) 4 16

Stony corals (Order Scleractinia) 57 77

Zoanthid gold coral (identified as Gerardia sp.) 1 0*

Gorgonian corals (Order Alcyonacea – in part

[formerly Order Gorgonacea])

30 68

True soft corals and stoloniferans (Order

Alcyonacea)

12 15

Sea pens (Order Pennatulacea) 1 10

Class Hydrozoa

Stylasterid corals (Order Anthoathecata,

Family Stylasteridae)

7 15

Total 112 201

characterize the benthic habitat and fauna of

the SAFMC’s new Snapper‐Grouper MPAs

from North Carolina to Florida (Harter et al.

2015; Figure 1; Appendix Table A). A total of

122 ROV dives surveyed 35 sites at depths

between 50‐200 m. Multibeam sonar surveys

mapped 574 km2 of benthic habitats. Other

studies in the region that shed light on deep‐sea

coral and sponge resources during this period

included exploration supported by NOAA’s

Office of Ocean Exploration and Research,

Harbor Branch Oceanographic Institute at

Florida Atlantic University, a Dutch/U.S.‐led

cruise on the R/V Pelagia (Mienis et al. 2014),

and surveys associated with proposed pipeline,

cables and offshore energy projects that

provided additional information on benthic

habitats of the Miami Terrace region (Messing

et al. 2012). Additional mapping of deep‐sea

habitats was conducted by the NOAA Ship

Okeanos Explorer and the U.S. Navy.

II.2. Taxonomy and Species

Distributions

II.2.i – Corals

a. Coral taxonomy

Ross and Nizinski (2007) reported 112 species

of corals from seven cnidarian orders from the

U.S. Southeast region (Table 1). Since 2007, the

systematic compilation of coral records in the

National Coral and Sponge Database, along

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Figure 2. Important structure‐forming deep‐sea corals of the Southeast U.S. a) Lophelia pertusa; b) ivory tree

coral (Oculina varicosa); c) large paramuriceid gorgonian coral with squat lobster and brittle stars; d) bamboo

coral (Keratoisis flexibilis); e) black coral (Leiopathes sp.); f) Stylasterid coral.

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b. Coral distribution

               

         

         

                 

                 

               

             

               

         

         

           

               

       

               

           

             

           

               

           

               

               

             

         

           

         

       

             

      

         

         

           

           

             

       

             

           

               

           

           

           

  

STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

with new descriptions and range extensions,

has nearly doubled this number to 201 (Table 1;

to relatively large sizes and occurring in

aggregations (Figure 2).

Online Annex 1; Cairns et al., this report). The

biggest increase in numbers of species was

among the gorgonian corals, reflecting reviews

by Cairns and Bayer (2009), DeVictor and

Morton (2010), and Watling et al. (2011). Calder

and Cairns (2009) described several new

stylasterid species from the region.

Research over the last ten years has expanded

our understanding of the distribution of

deepwater scleractinian coral habitats in the

region. Partyka et al. (2007), Harter et al. (2009),

Ross (2012), Reed et al. (2013a, 2014, 2015) and

David et al. (in prep.) summarized results of

numerous deepwater surveys from 2001 – 2011.

Ross et al. (2012) reviewed museum records of

dominant scleractinian corals from the region

and evaluated their utility for documenting

distributions. They cautioned that often such

records are misleading and in many cases are

incorrect. NOAA’s Database for Deep‐Sea

Corals and Sponges (Hourigan et al. 2015) has

instituted protocols to address these limitations

when possible. In addition, taxonomic and field

surveys of the shallower hard‐bottom habitats

(50‐150 m; DeVictor and Morton 2010, Harter et

al. 2015) have provided information on

components of the coral fauna that were not

addressed by Ross and Nizinski (2007). In both

the shallower and deep‐sea habitats, a number

of deepwater coral species, including reef‐

building stony corals, black corals, gorgonian

corals and stylasterid corals represent

important structure‐forming species – growing

Most deep‐sea corals (aside from

pennatulaceans) and sponges require hard

substrate for attachment. Thus the underlying

geology in the region determines the

distribution of such habitat, and along with

oceanographic conditions (especially those

generated by the Gulf Stream), influences the

distribution of deep‐sea corals (Ross and

Nizinski 2007, Reed et al. 2013a). Ross and

Nizinski (2007), Reed et al. (2013a) and others

have identified the following major deepwater

concentrations of hard‐bottoms in the Southeast

U.S. that support coral and sponge

communities:

1) Shelf and shelf break:  The continental shelf is

wide and shallow through most of the

region from Cape Hatteras to central

Florida. Hard‐bottoms are most common

near the shelf break (~ 55‐110 m depths) and

below the shelf break (110‐183m; see

SEAMAP‐SA 2001).  

2) Oculina coral mounds:  Off Florida, near the

shelf edge at depths of 70 – 100 m, are large

bioherms (10‐30 m relief) built by the

azooxanthellate form of the ivory tree coral,

Oculina varicosa (Reed 1980, 2002a,b, Reed et

al. 2007, Ross and Nizinski 2007, Maness

2011).   

3) Continental slope and Blake Plateau:  This is a

complex region with numerous hard‐

bottom and coral mound features

(lithoherms and bioherms). The continental

slope around Cape Hatteras is incised with

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

several canyons. South of Cape Hatteras,

the continental slope broadens from central

North Carolina into the Blake Plateau, a

broad depositional feature formed by the

Gulf Stream (400‐1250 m depths, Popenoe

and Manheim 2001). Major coral habitats in

this region include the North Carolina

Lophelia mounds, the Charleston Bump,

Stetson and Savannah banks, and extensive

Lophelia pertusa and Enallopsammia profunda

mounds off eastern Florida.

4) Miami and Pourtalès Terraces and

Escarpments: The Miami and Pourtalès

terraces are large limestone features

bordering the Florida Straits that provide

hard‐bottom habitat for deep‐sea corals and

sponges.

Shelf and shelf break: High relief ridges and

rock outcrops at the shelf break and on the

upper slope are often heavily encrusted with

sponges and gorgonians. These extensive hard‐

bottom habitats are also recognized as essential

fish habitat (EFH) for the snapper, grouper,

porgy species complex. In 2009, the South

Atlantic Council established seven deepwater

marine protected areas (MPAs) to help

maintain these stocks (see below and

http://safmc.net/managed‐areas/marine‐

protected‐areas; an eighth MPA, the Charleston

Deep Reef MPA is primarily sandy bottoms,

but has two large artificial reef sites).

Recent surveys of shelf‐edge MPAs by NOAA

have resulted in a better understanding of the

numerically dominant corals species present

(Harter et al. 2015). A total of 31 species of

corals were recorded from these habitats. These

were dominated by gorgonians (including,

Bebryce sp., Diodogorgia sp., Ellisella spp.,

Iciligorgia schrammi, Leptogorgia sp., Muricea sp.,

Nicella sp., Plumarella sp., Swiftia exerta, and

Titanideum frauenfeldii), which can reach

moderate sizes and occur in locally high‐

density patches (Fig. 3a). Several

azooxanthellate colonial stony corals also

occurred (Oculina varicosa, Madracis myriaster,

Madrepora oculata), but no bioherm formation

has been observed aside from the Oculina

bioherms off central Florida (see below). Other

coral species observed include black corals

(Antipathes atlantica, Antipathes spp., Stichopathes

sp., Tanacetipathes hirta), and soft corals

(including Anthomastus sp., Chironephthya

caribaea, Nidalia occidentalis and Telesto sp.).

Oculina coral mounds: The Oculina reefs

(bioherms) off eastern Florida are unique coral

habitats (Reed 1980, 2002a,b, Reed et al. 2013b).

These bioherms also serve as habitat for

gorgonians, soft corals, black corals and stony

cup corals (Fig. 3b,c). Recent work has

increased our understanding of the extent and

importance of these habitats. Harter et al. (2009)

reported on surveys of benthic habitat and fish

assemblages conducted inside and outside the

Oculina Bank Habitat Area of Particular

Concern (HAPC) in 2003 and 2005. They found

that the percentage of intact coral, fish

diversity, and densities of groupers were higher

inside the area closed to fishing.

In 2011, multibeam sonar and ROV surveys

discovered additional, previously unknown

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Figure 3. a) Shelf‐edge hard‐bottom habitat with Plumarella spp. octocoral colonies; b) supermale scamp grouper

(grey head color phase) on Oculina bioherm; c) large Oculina varicosa colonies.

Oculina coral mounds with live O. varicosa

stretching north of the then boundaries of the

Oculina Bank HAPC (Reed et al. 2013b). In

addition to the mounds, surveyed sites also

included smaller coral thickets, isolated live

colonies, and coral rubble on hard‐bottom.

These newly‐discovered and unprotected coral

mounds were similar in size to those within the

Oculina Bank HAPC; however, these mounds

had only 4.4‐5.5% cover of standing coral

framework, compared to 28.6‐37.8% cover

within the HAPC (Reed et al. 2013b). Most live

O. varicosa colonies at these unprotected sites

were much smaller in size compared to those

within the HAPC, perhaps reflecting recovery

following previous damage. Densities of live O.

varicosa and gorgonian colonies, sponges, and

fishes were also higher in the protected HAPC.

While there is some evidence that the Oculina

Bank HAPC has been effective in protecting

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

coral habitat, benthic biota, and fish populations

in recent years (Harter et al. 2009), coral and fish

densities remain severely depleted from what

they used to be in the 1980’s prior to the reefs

being decimated by trawling gear.

Oculina varicosa occurs as both shallow‐water

zooxanthellate and deepwater azooxanthellate

forms. Only the deepwater azooxanthellate

corals form bioherms. Eytan et al. (2009)

provided evidence from mitochondrial DNA

that O. varicosa corals from the Oculina Banks

were genetically isolated from the shallow‐

water zooxanthellate populations that occur

from North Carolina to the eastern Gulf of

Mexico. Genetic population separation by depth

has also been found in gorgonians in the Gulf of

Mexico (see Morrison et al. and Boland et al.,

this volume).

Continental slope and Blake Plateau: The

dominant framework‐forming scleractinian

corals in waters deeper than 200m are Lophelia

pertusa and, to a lesser extent Enallopsammia

profunda. Both species form deepwater coral

mounds or reefs, including bioherms and

consolidated limestone lithoherms, which are a

dominant feature on the Blake Plateau from

North Carolina to south Florida and the

Bahamas (Brooke 2007, Ross and Nizinski 2007 ‐

and references therein; Grasmueck et al. 2007,

Ross and Quattrini 2007, Messing et al. 2008,

Correa et al. 2011, 2012, Reed et al. 2013a).

Madrepora oculata occurs commonly in smaller

colonies, but does not appear to contribute

substantively to the reef framework. Partyka et

al. (2007) and Ross (2012) summarized habitat

information from submersible and ROV dives

from North Carolina to southeast Florida. Reed

et al. (2013a) compiled information on over 400

sites at depths between 200 and 900 m with

high‐relief bathymetry, from northeastern

Florida (31oN) through the Straits of Florida, of

which 147 sites were groundtruthed with

submersible or ROV dives. Ross et al. (2012)

analyzed museum specimens of four species of

deepwater branching stony corals (L. pertusa, E.

profunda, E. rostrata and M. oculata) from the

region.

The Cape Lookout and Cape Fear Lophelia

mounds off North Carolina appear to be the

northernmost bioherms in the U.S. Atlantic

(Reed et al. 2006, Ross and Nizinski 2007,

Partyka et al. 2007, Roberts et al. 2009). These

are extensive high profile (up to 80‐100 m tall)

bioherms almost entirely composed of dead L.

pertusa, entrapped sediments, and capped with

living and dead L. pertusa (Ross and Quattrini

2009, Mienis et al. 2014). These bioherms have

less cover of Enallopsammia spp. than do the

bioherms off the east coast of Florida. The North

Carolina mounds also appear to have fewer

black corals and octocorals compared with

regions to the south and the Gulf of Mexico.

Mienis et al. (2014) deployed landers to measure

near‐bottom environmental conditions on coral

mounds near Cape Lookout. They found that

the corals were exposed to large temperature

and current fluctuations related to Gulf Stream

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Figure 4. Lophelia coral bioherms: a) Cape Lookout Lophelia pertusa habitat; b) dense live Lophelia pertusa

off E. Florida; c) codling (Laemonema melanurum) on coral habitat; d) Enallopsammia profunda coral and a

hexactinellid sponge (Aphrocallistes beatrix).

           

             

             

               

       

               

             

             

           

         

            

             

                

             

             

             

           

             

               

       

           

                 

               

             

                 

             

STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

meanders, and concluded that L. pertusa live

under extreme conditions that may limit present

mound growth. However, in the same area

Brooke et al. (2013) noted that L. pertusa survived

these extreme and variable temperatures,

indicating resilience in this species. Matos et al.

(2015) investigated the age of the Cape Lookout

coral mounds using cores and U‐series dating

and concluded that mound growth primarily

occurred during interglacial periods, most

recently during the last 7,000 years.

In contrast to North Carolina, southern Georgia

and Florida, coral habitats of the central Blake

Plateau region appear to have mostly smaller

aggregations of live corals occurring both on

coral mounds and attached to abundant rocky

substrata. Off South Carolina and Georgia,

Stetson and Savannah banks and the Charleston

Bump represent major areas of high relief, hard

bottom topography with significant coral

habitats, including Lophelia mounds (Partyka et

al. 2007, Ross and Quattrini 2009, Sedberry et al.

2010). The Charleston Bump rises from 700m to

400m and has a varied topography ranging

from flat hard bottom to rugged relief and near

vertical scarps (Sedberry 2001). Corals in the

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

area include small colonies of Lophelia pertusa,

Madrepora oculata, and Enallopsammia profunda;

various stylasterids, black corals and

gorgonians (Partyka et al. 2007).

At Stetson Banks on the Blake Plateau, Reed et

al. (2006) identified of 11 coral taxa. The

colonial Scleractinia were dominated by

colonies of L. pertusa (30–60 cm tall), along with

E. profunda and Solenosmilia variabilis. Small

stylasterid corals were common and numerous

species of solitary cup corals were abundant.

They reported at least six taxa of octocorals,

dominated by primnoids (Family Primnoidae),

Paramuricea spp. (Family Plexauridae), bamboo

corals (Family Isididae), as well as smaller

stoloniferan and nephtheid soft corals.

The Savannah Banks off Georgia include

numerous lithoherms at depths of 490 to 550 m

along the western Blake Plateau. Corals and

sponges occurred on exposed terrace

pavements, with the greatest abundances along

the rock outcrop edges and on pinnacle crests

(Reed et al. 2006, Partyka et al. 2007). The

estimated cover of sponges and gorgonians was

10% on exposed rock areas. Colonies of L.

pertusa (15–30 cm diameter) were common but

only accounted for ~1% of the benthic cover and

were intermixed with small colonies of M.

oculata and stylasterid hydrocorals. Primnoids,

plexaurids and black corals were also observed.

Diversity of octocorals, black corals and

sponges increases from this point south on the

Blake Plateau, and the branching stony corals

Madrepora oculata and Enallopsammia profunda

become more abundant.

Lophelia pertusa and E. profunda coral mounds

appear to reach their highest densities from the

Jacksonville Lithoherms off southern Georgia

and N. Florida, and stretching southwards

through the Straits of Florida (Reed et al. 2013a;

Fig. 5). New multibeam sonar mapping surveys

conducted since 2007 have revealed the

abundance of the mound‐structures,

particularly off eastern Florida. Reed et al.

(2013a) estimated that this habitat off eastern

Florida covers over 13,400 km2. Live coral cover

ranges from less than 5% to nearly 100% on the

peaks and up‐current slopes facing the Gulf

Stream or Florida Current, a component of the

Gulf Stream that flows through the Florida

Straits separating Florida from Cuba and the

Bahamas (Reed et al. 2013a). These mounds are

predominately L. pertusa but some are

apparently entirely E. profunda, and some are

intermixed. Madrepora oculata is not a major

component of these mounds and occurs only as

isolated, small colonies with either L. pertusa or

E. profunda. For reasons unknown, extensive

areas of dead E. profunda rubble are also

common throughout the region and often

observed at the bases of the mounds and the

foot of the Miami Terrace. The mounds also

provide substrate for other structure‐forming

taxa, such as gorgonian, black and stylasterid

corals and sponges.

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Figure 5. Coral regions of deep‐sea coral habitats mapped from northeastern Florida (31o

N) through the Straits of

Florida. Dashed gray line = U.S. EEZ; polygons (heavy bold line) = boundaries of deepwater Coral Habitat Areas

of Particular Concern (CHAPC) and deep‐water Oculina Bank HAPC; colored polygons = deep‐sea coral habitat

regions and major geological/topography features (see legend). Stars = Deep‐sea coral habitats mapped with ROV

or submersible. Museum records from National Museum of Natural History, Smithsonian Institution: Dots =

Lophelia pertusa, Enallopsammia profunda, Madrepora oculata or other deep‐water coral species (> 50 m

depth). See Reed et al. (2013c) for methodology used to identify the deep‐sea coral habitat regions (Data from Reed

et al. 2005, 2006, 2013c, 2014). Depth contours in meters.

STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

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Figure 6. Miami Terrace: a) Rock pavement and ledges provide habitat for a variety of species including black

corals, stylasterid corals, gorgonians, and sponges; b) diverse habitat of sponges, stylasterid corals, pencil urchins

and squat lobsters; Pourtalès Terrace: c) Ledge with dense cover of stylasterid corals; d) Deepwater Lophelia

pertusa coral mound; live coral colony (~1 m diameter), black coral (Leiopathes sp.), and mora codling

(Laemonema melanurum).

         

         

         

                 

           

                 

                   

           

               

             

             

       

             

           

             

           

         

           

               

             

           

   

STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Several relatively shallow bioherms have

recently been identified from multibeam

surveys off northeastern Florida. Habitat

surveys in 2010 revealed a matrix of L. pertusa,

including living colonies fused into thickets

along with dead colonies and rubble at the crest

of these bioherms at around 210 m (Ross et al.

2015a). This is significantly shallower than

previous records of live L. pertusa in the region,

which generally occurs from around 360‐800 m

(Ross and Nizinski 2007). These represent the

shallowest recorded bioherms with large living

colonies from the western North Atlantic (Ross

et al. 2015a). Other corals (e.g., Leiopathes sp.)

and fishes commonly observed on deeper reefs

were also observed in these sites.

Measurements of environmental parameters at

these sites revealed colder, nutrient‐rich waters,

leading Ross et al. (2015a) to conclude that

these sites were subject to persistent conditions

of Gulf Stream‐driven upwelling of deep,

nutrient‐rich water.

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Miami and Pourtalès Terraces: The upper

Miami Terrace extends for 145 km off southeast

Florida at around 275 m depth (Brooke 2007,

Grasmueck et al. 2007a,b, Messing et al. 2008,

Vinick et al. 2012, Reed et al. 2006, 2013a). The

bordering escarpment drops steeply to a lower

terrace at depths of 500 – 700 m, with low

discontinuous ridges perpendicular to the

current and patchy colonies of L. pertusa and E.

profunda, as well as gorgonians, stylasterids,

and sponges (Fig. 6 a,b). The slope below the

lower terrace (710‐870 m) has a series of much

longer ridges reaching 20 m in height and

capped with sponges and various corals, L.

pertusa, stylasterids, isidids and other octocorals

(Correa et al. 2012). Enallopsammia coral

mounds and coral rubble are the dominant

habitat types near the foot of the escarpment.

Pourtalès Terrace stretches 213 km at depths of

200 – 450m along the southern edge of the Flor‐

ida Keys reef track and provides extensive, high

relief, hard‐bottom habitat (Gomberg 1976,

Reed et al. 2005, 2013c, 2014). Hard‐bottom,

topographic features consisting of numerous

high‐relief mounds and ridges on the mid‐

terrace and a chain of sinkholes extend for ~100

km along the southwest terrace margin.

Stylasterids appear far more abundant on

Pourtalès Terrace, compared to the nearby

Miami Terrace. Lophelia and Enallopsammia

mounds appear to be relatively rare, though

surveys in 2011 revealed a Lophelia mound, the

southernmost mound known from U.S. waters

(Reed et al. 2014; Fig. 6 c,d). The reasons for

these faunal differences between the Miami and

Pourtalès terraces remain unclear. Cairns (1992)

hypothesized that distributions of stylasterids

may be partially limited by their preference for

oligotrophic waters and sensitivity to

fluctuating salinity and sedimentation often

associated with continental margins.

Broad‐scale biogeographic distribution

patterns: Prior to 2007, most distributional

studies focused on stony corals (e.g., Cairns and

Chapman 2001), with particular interest in the

framework‐forming corals such as L. pertusa,

and O varicosa (Ross and Nizinski 2007). While

there was recognition that gorgonian and black

corals were important components of habitat

(Ross and Nizinski 2007), this report includes

the first regional maps of gorgonian and black

coral distributions (see appendix maps). A

number of species of gorgonian and black

corals are associated with the Lophelia coral

mounds, but they are not limited to these

habitats. For example, gorgonians (e.g.,

Plumarella sp., Swiftia exserta) can reach high

densities in shelf‐edge habitats. Black corals

and hexactinellid sponges are relatively rare on

the North Carolina coral banks (Ross and

Quattrini 2009), but are common further south

(Reed et al. 2006).

The distribution of stylasterid corals also

suggests discontinuities within the Southeast

U.S. region. These species are relatively small,

but colonies can occur in dense aggregations in

the Caribbean and the southern portion of the

region, being especially important on Pourtalès

Terrace (Reed et al. 2014). They are largely

absent north of 32.5o N latitude.

The Florida Current appears to form a

biogeographic barrier for a number of coral

species. Numerous coral species that are

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a. Sponge taxonomy

               

             

                 

             

             

               

               

               

       

         

           

             

           

             

               

STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

common in the Bahamas and the Caribbean

appear to be rare or absent in U.S. waters off

Florida. For example, the zoanthid gold coral,

Kulamanamana sp. (formerly known as Gerardia

sp.) occurs in dense aggregations along

upcurrent crests of mounds at depths of 500 –

700 m in the Bahamas (Messing et al. 1990), but

this taxon has not been recorded from the U.S.

side of the Florida Straits (C. Messing, pers.

comm.). There are several other examples of

coral species that have not been reported from

southeast Florida waters, but occur nearby

across the Florida Straits. These include two

black corals (Distichopathes filix and

Tanacetipathes tanacetum) and a stony coral

(Balanophyllia (Balanophyllia) cyathoides) both

found in Cuba and the Bahamas, and two

additional stony corals (Caryophyllia

(Caryophyllia) antillarum and Deltocyathus

agassizi) and a soft coral (Chironephthya agassizii

= Siphonogorgia agassizii) known from the

Bahamas. A single species of stony coral

(Thalamophyllia gombergi) appears to be endemic

to the Florida side of the straits and has not

been reported from Cuba or the Bahamas

(Cairns and Chapman 2001).

Morrison et al. (2011) used microsatellite DNA

markers to examine population connectivity in

L. pertusa in the North Atlantic, including

samples from the Southeast U.S. They found

distinct genetic groupings among corals

collected in the Gulf of Mexico, Southeast U.S.,

New England Seamounts, and Northeast

Atlantic, suggesting that these represent

distinct populations. The corals from the

Southeast U.S. were distinguishable from, but

most closely related to those from the Gulf of

Mexico. The Gulf of Mexico Loop Current

draws warm water from the Caribbean into the

eastern Gulf, then loops south along the west

Florida slope, feeding into the Florida Current

through the Florida Straits, and then into the

Gulf Stream. This current system likely

transports pelagic larvae from the eastern Gulf

of Mexico to the Southeast region (Morrison et

al. 2011). Hare and Walsh (2007) used satellite‐

tracked drifters released from three MPAs (the

Tortugas South Ecological Reserve, Oculina

Bank HAPC and Gray’s Reef National Marine

Sanctuary) along with transport models to

investigate larval transport in the region. The

results predicted export of larvae by the Florida

Current/Gulf Stream, but with some retention

in the lower Florida Keys and off Georgia.

II.2.ii – Sponges

Unlike deep‐sea corals, there has not been a

systematic inventory of sponge taxa from deep

waters of the Southeast Region. Van Soest et al.

(2012) reported nearly 300 described species of

shallow and deepwater sponges in the three

ecoregions that are in or adjacent to the

Southeast U.S. (Carolinean – 34, Floridian – 153,

and Bahamian – 95) (Table 2). These are

predominantly in the Class Demospongiae,

however the authors acknowledge that deep‐

sea sponge species are under‐reported. Most

sampling in the region has focused on shallow‐

water, easily accessible species. The Class

Hexactinellida (or glass sponges) is found at

depths greater than 200m, and along with the

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Figure 7. Important structure‐forming deep‐sea sponges of the Southeast U.S. Demosponges: a) Discodermia

sp.; b) Spongosorites sp.; c) newly described species Theonella wrightae Pisera and Pomponi, 2015;

Hexactinellid glass sponges: d) Rhadopectella tintinnus; e) Aphrocallistes beatrix; f) Nodastrella

asconemaoida from Miami Terrace.

             

             

             

               

             

           

               

     

             

           

           

           

   

   

             

               

             

         

                 

         

               

               

                 

        

           

               

               

       

           

             

             

             

STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

perception that this group is difficult to

identify, the number of described species is

low, relative to the Class Demospongiae (Table

2). Species within the Class Calcarea occur in

both shallow and deep water, but also

considered difficult to identify. Few experts

have focused on these two groups. The Class

Homoscleromorpha is likely under‐reported

because many of the deepwater species are

thinly encrusting and difficult to collect.

An example of unexplored and largely

undocumented deep‐sea sponge fauna are the

carnivorous sponges (Demospongiae:

Poecilosclerida: Cladorhizidae). This unusual

group of sponges differs significantly in body

plan and cellular organization from the rest of

the Phylum Porifera, and was only recently

discovered (Vacelet and Boury‐Esnault 1995).

Since that time, at least 41 new species of

carnivorous sponges have been described,

primarily in deep water (Cristobo et al. 2015).

Nine species are found in the Caribbean region,

four of which occur off the coast of Florida

(Hestetun et al., 2016).

Several recent publications (e.g. Vacelet 2006,

Vacelet et al. 2009, Kelly and Vacelet 2011,

Lopes and Hajdu 2014, Hestetun et al. 2017)

have demonstrated that identification of

sponges from previously poorly known areas

has resulted in a high diversity of carnivorous

sponge species. It is likely that further

examination of the sponge fauna from the

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b. Sponge distribution

           

             

           

       

         

         

         

         

               

       

       

           

             

               

       

           

               

           

             

               

             

‐ ‐

STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Table 2. Number of shallow and deepwater sponge species reported by van Soest et al. (2012) from the

Carolinean, Floridian, and Bahamian ecoregions (which encompass the Southeast U.S. and neighboring areas)

compared with the number of deepwater sponge taxa reported in this chapter (from Reed et al. 2005, 2006,

2013c, 2014, 2015). Note that deep‐sea sponges are under‐represented in the numbers reported by van Soest et

al. (2012) and are only used for comparison with the numbers of deep‐sea sponges reported in this chapter.

Taxon Van Soest et al. (2012)

(primarily shallow water)

This report

(50 816 m)

Class Demospongiae 267 98

Class Hexactinellida 7 18

Class Homoscleromorpha 1 5

Class Calcarea 7 7

Total 282 121

Southeast U.S. will yield additional species, not

only in the Cladorhizidae, but also in the

Hexactinellida, Homoscleromorpha, and the

Calcarea.

A number of sponges have been collected

under an ongoing program of biomedical and

biotechnology research at Harbor Branch

Oceanographic Institute, Florida Atlantic

University. This program has provided

museum voucher specimens and information

on taxonomy, morphology and chemistry of

species and the habitats in which they are

found. More recent expeditions by Harbor

Branch and NOAA have focused on

quantifying density and diversity of benthic

invertebrates using photographic surveys.

While some common taxa can be identified to

genus or species, many can only be identified to

a higher level such as class, order, or family.

Sponges are especially difficult to identify

without a sample that can be used for

microscopic analyses of the skeleton.

Within the Class Demospongiae, the most

common deep‐sea taxon that occurs in the

Southeast U.S. region is the Order

Tetractinellida (formerly Astrophorida). In

general, worldwide, this order includes massive

sponges that dominate hard‐bottom deep‐sea

ecosystems, both in biomass and abundance.

Additionally, the Tetractinellida also includes

the “lithistid” or rock sponges, members of a

polyphyletic group, which occur worldwide,

primarily in deep‐water environments of

temperate and tropical latitudes (Maldonado et

al. 2016). Lithistid sponges are characterized by

spicules that have become fused into a rigid

silica framework. Recently, biological and

chemical interest in lithistid sponges has

increased, since these sponges are the source of

more than 300 novel and diverse compounds,

many of which have potential human health

applications (for review, see Winder et al. 2011).

Pomponi et al. (2001) described the diversity

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Figure 8. The lithistid demosponge, Leiodermatium sp. (foliose tan sponge in the middle), is an important

structure‐forming deep‐sea sponge on the Miami Terrace. It is also the source of potent anticancer compounds.

and bathymetric distribution of lithistids

collected during expeditions to the Southeast

U.S. Thirty species of lithistid sponges have been

reported from the tropical western Atlantic

region (which includes the U.S. Southeast

Region), but unpublished data suggest a much

higher number of species present in this region

(Pisera and Pomponi 2015).

Certain tetractinellid sponges form aggregations,

known as “sponge grounds” which have been

best documented further north in the Atlantic

(e.g., northeastern Atlantic, Klitgaard and

Tendal 2004; northwestern Atlantic. Knudby et

al. 2013). Aggregations have been observed in

the Southeast U.S. and Caribbean (e.g.,

Maldonado and Young, 1996), but are much

smaller in extent. These sponges, along with the

glass sponges (Order Hexactinellida), reach the

size and density to suggest that they have

particular significance related to provision of

habitat, bentho‐pelagic coupling, and

biogeochemical cycling (reviewed by

Maldonado et al. 2016).

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

Sponges are important components of the

sessile epifauna and occur in most of the same

habitats that harbor deep‐sea corals. The

descriptions below summarize observations on

sponge distributions in the same major habitat

regions previously reviewed for corals.

Shelf and shelf break: Sponges are an important

component of the benthic macrobiota in hard‐

bottom habitats along the continental shelf

break. Harter et al. (2015) reported 52 taxa of

sponges from surveys of the South Atlantic

MPAs established under the Council’s Snapper‐

Grouper Fishery Management Plan. Sponge

cover ranged from < 1% to over 6%. The

dominant sponges were demosponges in the

Orders Agelasida (Agelas sp.), Axinellida

(Auletta sp., Scopalina sp.), Biemnida

(Neofibularia nolitangere), Chondrillida

(Chondrilla sp.), Chondrosiida (Chondrosia sp.),

Clionaida (Spirastrellidae, Cliona sp.);

Dictyoceratida (Ircinia campana, Ircinia strobilina)

Haplosclerida (Callyspongia vaginalis, Niphates

sp., Oceanapia sp., Siphonodictyon coralliphagum,

Xestospongia muta), Poecilosclerida (Clathria sp.,

Desmapsamma anchorata, Zyzzya sp.),

Polymastiida (Polymastia sp.), Suberitida

(Spongosorites sp.), Tetractinellida

(Cinachyra/Cinachyrella sp., Corallistidae, Erylus

sp., Geodia sp., Leiodermatium sp., Theonella sp., ),

and Verongiida (Aiolochroia crassa, Aplysina sp.).

Only one species (Farrea sp.) of Hexactinellida

(glass sponge) was identified.

Oculina Coral Mounds: Eighteen species of

sponges have been identified from the deep‐

water Oculina mounds. Several species of

Geodia, Eylus, Ircinia, and unidentified demo‐

sponges dominated the sponge assemblage

(Reed et al. 2013b, Harter et al. 2015).

Continental slope and Blake Plateau: Occurring

principally in areas of hard substrate (e.g., coral

mounds, rocks and ledges, etc.), sponges are

among the important and obvious components

of epifaunal communities in the deeper water

areas surveyed on the continental slope and

Blake Plateau (Partyka et al. 2007, Reed 2002b,

Reed et al. 2006, Ross 2010). The sponge fauna

includes both demosponges (Class

Demospongiae) and glass sponges (Class

Hexactinellida), many of which can reach much

larger sizes than generally found on the

shallower Oculina mounds (Reed 2002a,b). As

with corals, the diversity of sponges appears to

increase south of Cape Fear (Partyka et al. 2007).

Reed et al. (2006) identified 18 sponge taxa on

the Stetson Banks. The most abundant taxa

were demosponges (Geodia spp., Leiodermatium

spp., and sponges in the families

Pachastrellidae and Corallistidae) and glass

sponges. On the steeper, upper flanks of the

mounds (625 to 671 m), the density, diversity,

and size of sponges increased, with 15–50 cm

sponges most abundant. Massive Spongosorites

spp. and hexactinellids were common, and

pachastrellid tube sponges were abundant. On

the top of plateaus, the dominant sponges were

fan sponges (Phakellia spp.) and numerous other

demosponges.

On the Savannah Lithoherms, 12 sponge taxa

were identified, occurring principally on

exposed terrace pavements, with greatest

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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abundance observed along the edges of rock

outcrops and on pinnacle crests. Dominant

sponges included large fan sponges (Phakellia

ventilabrum), plate sponges (pachastrellids and

tetractinellids), and hexactinellid glass sponges.

Some ridges and terraces at some sites had

nearly 100% cover of sponges (Phakellia spp.,

Geodia spp., Pachastrellidae, and hexactinellids;

Reed et al. 2006)

On the East Florida Lophelia mounds, Reed et al.

(2006) identified 18 sponge taxa. Dominant

sponges consisted of numerous demosponges

and dominated by Phakellia spp., Geodia spp.,

Spongosorites spp., pachastrellids and petrosiids.

Hexactinellids were also common, especially

Aphrocallistes beatrix and species in the

family Euretidae.

Miami and Pourtalès Terraces and

Escarpments: The Miami Terrace and

Escarpment host diverse and dense sponge

assemblages (Reed et al. 2006), including

hexactinellids (e.g., Heterotella sp.) and

demosponges (including Spongosorites spp.,

Geodia spp., Vetulina spp., Leiodermatium spp.,

Petrosia spp., and sponges in the families

Raspailiidae, Tetractinellida, Pachastrellidae,

and Corallistidae). In separate studies on

Pourtalès Terrace, Reed et al. (2005, 2014)

identified 91 sponge taxa from the quantitative

ROV and submersible photo‐transects. Of these

taxa, 48 were identified to genus or species

level, and four are possible new species.

Thirteen sponge taxa were found solely in the

sinkholes and not on the rock mounds. A newly

described species, Phakellia wrightae (Pisera and

Pomponi 2015), and other sponges in the

Family Theonellidae were found exclusively on

the mounds. Other dominant taxa included

Geodia spp. (three species new to science [Paco

Cardenas, Evolutionary Biology Center,

Uppsala University, Sweden, personal

communication]), Pachastrella sp., Corallistidae,

Theonella spp., Leiodermatium sp.; encrusting

Spirastrellidae; Poecilosclerida (Raspailliidae,

encrusting Hymedesmia sp.); Auletta sp.;

Topsentia sp.; and Homoscleromorpha (Plakortis

spp.). Glass sponges included Aphrocallistes

beatrix beatrix, Hexactinella sp., Iphiteon sp., and a

recently described genus, Nodastrella

(Dohrmann et al. 2012). More surveys were

conducted on Pourtalès Terrace than in other

regions, which may explain in part the higher

number of sponges identified. Densities of

sponges were very high, especially on the

plateaus and terraces of the mounds on

Pourtalès Terrace. Maximum densities of

sponges (> 5 cm) on the plateaus ranged from 1

to 80 individuals m−2. The walls of the sinkholes

were also densely encrusted with sponges (1 to

50 individuals m−2).

Broad‐scale biogeographic patterns:

Information on the distribution of sponges is

currently too fragmentary to draw many

conclusions about their geographic distribution

patterns. There are distinct differences between

the shelf‐edge (< 150 m) sponge assemblages

and those in deeper habitats. The former are

dominated by demosponges (e.g., Ircinia spp.,

spirastrellids), while the latter are characterized

by more demosponges of Order Tetractinellida

(e.g., pachastrellids, corallistids, geodiids) and

Axinellida (e.g., Phakellia spp.), along with

significant numbers of glass sponges.

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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II.3. Species Associations with

Deep-Sea Corals and Sponges

Ross and Nizinski (2007) summarized

information on species associations with deep‐

sea coral habitats. Since then, there have been

additional studies involving deep‐coral and

faunal associations within this region. In many

cases, sponges are also an important contributor

to 3‐dimensional structure in the same habitats

where corals are present, but this habitat

component remains poorly documented.

Many deep‐sea fishes and perhaps other fauna

in the North Atlantic Ocean are closely

associated with complex habitats, at least on the

upper slope (200‐1000 m). However, the specific

roles of deep‐sea corals in structuring or

influencing communities remain unclear (Ross

et al. 2015b). Although there are a few studies

from other regions indicating that certain fishes

or invertebrates may use corals for specific

activities (spawning, nurseries, feeding; e.g.,

Buhl‐Mortensen and Mortensen 2004, Mah et al.

2010, Baillon et al. 2012, Henry et al. 2013, Reed

et al. 2014, Ross et al. 2015b), most studies have

not demonstrated a statistically significant link

in faunal occurrence to living corals or

demonstrated a functional relationship for

fishes (Auster 2007, Ross and Quattrini 2007,

Harter et al. 2009, Biber 2014, Ross et al. 2015b).

Often standing dead coral framework has

proved to be as significant a habitat for

associated fauna as living coral (Lessard‐Pilon

et al. 2010, Quattrini et al. 2012). It is important

to stress that fauna may be mostly influenced

by habitat structure (size, profile, rugosity, etc.)

and not any one component of the habitat (e.g.,

corals). However, deep‐sea corals are a major

contributor to overall habitat complexity on the

slope and in some areas are the dominant

habitat component (as in coral‐built bioherms).

Quattrini et al. (2015) indicated that coral

diversity was important in explaining variation

in fish assemblages in the Northeast U.S.

Canyons region. Thus, considering that corals

are a major contributor to deep‐sea habitat

complexity and that fauna is often more diverse

and abundant in such habitats (as suggested by

Milligan et al. 2016), it seems most prudent to

consider deep‐sea corals as Essential Fish

Habitat (EFH) for associated fishes.

Shelf and shelf break: Shelf‐edge habitats are

home to 167 different fish species, including 33

species from the snapper‐grouper complex

(Quattrini and Ross 2006, Schobernd and,

Sedberry 2009, Harter et al. 2015). The

distribution of coral and live hard‐bottom

habitat in this region (to about 300 m) is

considered a proxy for the distribution of the

species within the snapper grouper complex

(SEAMAP‐SA 2001, SAFMC 2007a). Harter et

al. (2015) and Quattrini and Ross (2006) found

distinct differences in fish communities from

shelf‐edge habitats in different geographic

areas. Surveys were not able to discriminate the

extent to which the association of fishes might

have been influenced by the 3‐dimensional

structure provided by corals or sponges,

distinct from the topography of the rocky hard‐

bottoms themselves. Quattrini and Ross (2006)

noted that species richness was greatest in the

highest profile habitats. These shallower

habitats do not include coral bioherms, and the

relative role corals and sponges in providing

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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habitat structure for fishes compared with the

underlying rocky outcrops has not been teased‐

out. Corals and sponges are also important

habitat for numerous invertebrate species,

however we are not aware of any studies of

these associations for shelf and shelf break

hard‐bottom habitats in the Southeast U.S.

region.

Oculina Coral Mounds: The Oculina biogenic

refuge consists primarily of standing live and

dead coral habitat (Reed et al. 2007). As long as

the coral is standing, the living space within the

colony branches supports dense and diverse

communities of associated invertebrates (Reed

et al. 1982, 2002a,b, Reed and Mikkelsen, 1987).

However, once reduced to unconsolidated coral

rubble, little living space is left except for the

infauna (Reed, 1998). A hypothetical trophic

model of the Oculina varicosa ecosystem

indicates that a significant loss of habitat, in

particular intact live and dead standing coral,

could bring dramatic shifts in the ecosystem

(George et al. 2007). Unfortunately, such a loss

of habitat on the Oculina reefs has occurred, due

to extensive mechanical damage from bottom

trawling for rock shrimp (Sicyonia brevirostris)

(Reed et al. 2007). Between 1977 and 2001,

submersible surveys found nearly 100% loss of

live‐coral cover (range 98.4%–100%) outside of

the protected Oculina Bank HAPC boundaries,

whereas the only reefs that survived (e.g., –

Jeff’s and Chapman’s Reefs) were within the

protected area. Fish populations experienced

dramatic declines (Koenig et al. 2000, 2005).

Even within the Oculina Bank HAPC, several

reefs were destroyed (e.g., Sebastian Pinnacles,

Steeples) between 1977 and 2001, apparently as

a result of illegal fishing, highlighting the

importance of effective enforcement.

The importance of the standing coral for

biodiversity of the region is evident in the

associated community of invertebrates found in

the coral. Collections of 41 O. varicosa colonies

(~15‐20 cm diameter), which were broken up

and sieved through 0.5 mm mesh, resulted in

20,000 invertebrate specimens. These

collections, along with additional qualitative

collections by manned submersibles and

dredge samples of coral rubble, have

documented the following biodiversity within

the Oculina habitat: 230 species of mollusks, 50

decapods, 47 amphipods, 21 echinoderms, 15

pycnogonids, 100+ polychaetes. 42 bryozoans,

11 sipunculans, and numerous other taxa

(references in Reed et al. 2006, Winston 2016; M.

Rice, Curator emeritus, Smithsonian Natural

History Museum, pers. comm.).

Harter et al. (2009) analyzed fish assemblages

and benthic habitats inside and outside the

Oculina Experimental Closed Area (OECA), in

2003 and 2005 using ROVs. The OECA was

closed to all bottom fishing in 1994 in order to

evaluate the effectiveness of the reserve for the

management and conservation of managed fish

populations. A total of 62 fish species were

observed, including economically‐important

grouper species such as speckled hind

(Epinephelus drummondhayi), red grouper (E.

morio), snowy grouper (Hyporthodus niveatus),

scamp (Mycteroperca phenax), gag (M.

microlepis), and rock hind (E. adscensionis; not

typically a target species but kept as bycatch).

Even though fish assemblages were not

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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different among management areas,

biodiversity and grouper densities were higher

inside the OECA compared to outside. The

study did not track coral restoration, however,

the percentage of intact coral was also higher

inside the OECA. These results provide initial

evidence demonstrating effectiveness of the

OECA for restoring reef fish populations, and

add support to the value of intact O. varicosa

habitat for these species. Additionally, gag and

scamp were known to aggregate on and use O.

varicosa for spawning habitat and juvenile

speckled hind use the coral for shelter suggest‐

ing a nursery value of the coral (Gilmore and

Jones 1992, Koenig et al. 2000, 2005).

New ROV surveys provided some indication of

potential recovery by populations of certain fish

species. In the 1980s, large, mature black sea

bass (Centropristis striata) dominated the O.

varicosa reefs, but had disappeared by the 1990s

(Koenig et al., 2000). In 2005, black sea bass,

mostly small juveniles, ranging in length from

10 to 20 cm, were observed, suggesting initial

stages of recovery for this species. Subsequent

ROV surveys in 2015 and 2016 observed black

sea bass throughout the Oculina region,

including Chapmans Reef inside the OECA, in

the newly designated northern extension area

of the Oculina Bank HAPC, as well as in the

satellite areas of the Oculina Bank HAPC

(Harter, personal communication). All of these

findings combined present initial evidence

demonstrating effectiveness of the OECA for

restoring reef fish populations and their habitat.

Continental slope and Blake Plateau: Ross and

Quattrini (2009) analyzed fish assemblages at

nine Lophelia‐dominated sites at depths of 366‐

770 m along the Southeast U.S. slope between

Cape Lookout, North Carolina to Cape

Canaveral, Florida. They found regional

differences in the deep‐reef associated fauna,

with sites off North Carolina differing from

those off the central Blake Plateau and off Cape

Canaveral. Depth and habitat structure seemed

to be the drivers of these differences. Quattrini

et al. (2012) in a smaller scale study noted that

habitat structure, especially height off bottom,

influenced faunal composition.

Because these deep reefs are difficult to sample

and remain poorly studied, there continue to be

discoveries of new species and new

distributional records for the area resulting

from new surveys. Since 2007, four new species

of benthic fishes were described (McCosker and

Ross 2007, Fernholm and Quattrini 2008,

Nielsen et al. 2009). Caruso et al. (2007) and

Pietsch (2009) reviewed the lophiform fishes

from the Gulf of Mexico and Southeast U.S.

deep reefs, noting many large range extensions.

Similarly, Henry et al. (2008) reviewed the

deepwater hydroids collected on Lophelia and

rocky reefs and reported a number of species

new to the southeast region.

Reed and Farrington (2010) and Reed et al.

(2017) analyzed video from submersible and

ROV surveys to document the occurrence and

distribution of golden crab (Chaceon fenneri) in

deep‐water habitats (200‐900 m deep), including

coral habitats, off eastern and southern Florida.

Golden crabs were observed at depths of 247—

888 m, with most crabs found between 400‐

500m. While crabs were observed in habitats

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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ranging from dense live coral thickets to flat soft

bottoms, Reed et al. (2017) found that densities

were significantly higher on soft‐bottom habitat

(sand and mud) compared to hard‐bottom

habitat (rocky or coral‐dominated habitats).

Within the hard‐bottom habitats, mean crab

density appeared to be higher on rocky substrata

(pavement, rock slabs, boulders, cobble and

rubble) compared to coral substrata (standing

live/dead coral, coral rubble). The authors noted

that it may be easier to see crabs on open flat

soft‐bottom habitat compared to more complex

habitats, however, they found no evidence that

crabs were hiding or sleeping in crevices during

the daytime surveys.

Reed and Farrington (2010) also reported on

observations of three other economically

important fisheries species, blueline tilefish

(Caulolatilus microps), golden tilefish

(Lopholatilus chamaeleonticeps), and royal red

shrimp (Pleoticus robustus). Observations of

tilefishes were limited, although five out of

seven blueline tilefishes were observed

associated with high‐relief bioherms on the

Pourtalès Terrace (186 to 285 m). Royal red

shrimp are also an important species targeted

by fishers at these depths, but were difficult to

document using submersibles or ROVs.

Pourtalès Terrace: Reed et al. (2015) reported

over 50 fish species, including 11 economically‐

important species observed in deepwater

habitats of Pourtalès Terrace. Thirteen species

observed on Pourtalès Terrace (Reed et al. 2005,

2006, 2015) are also found on deepwater

Lophelia reefs from North Carolina to Cape

Canaveral (Ross and Quattrini 2007). Along the

Pourtalès Terrace and escarpment, deepwater

fish assemblages differed by depth (150‐300 m

and 450‐850 m), habitat geomorphology and

substrate (Reed et al. 2015). The fish

assemblages associated with Lophelia coral

habitat on Pourtalès Terrace appeared to differ

from assemblages on other pavement and rock

wall substrates, but the associations did not

appear to be obligate. Additionally,

Baumberger et al. (2010) reported a large

spawning aggregation of beardfish (Polymixia

lowei) associated with a sinkhole on the outer

edge of the Pourtalès Terrace.

Sponge associations: Sponges are known to

host significant communities of associated

invertebrates (Hogg et al. 2010). Sedberry et al.

(2004) analyzed five individual sponges of

different genera collected off South Carolina

and NE Florida at depths between 50‐194m.

They reported 947 invertebrates representing

ten taxonomic groups living in the sponges.

Polychaetes accounted for 69% of the faunal

associates. Amphipods (14%), cirripeds (8%),

and decapods (6%) were also important

constituents of the associated fauna. The

number and composition of faunal associates

appeared to vary by sponge species. Erylus sp.

hosted primarily annelids (96%); Chondrilla sp.

hosted primarily arthropods (77%), including

amphipods, decapods, and isopods; while

annelids and arthropods were both abundant

on Aplysina archeri and Ircinia campana

(Sedberry et al. 2004). Unfortunately, there are

no comparable studies on sponge communities

from deeper waters in this region.

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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II.4. Predictive Habitat Modeling of

Deep-Sea Corals

Only a very small portion of the deeper waters

off the Southeast U.S. has been surveyed for

deep‐sea coral or sponge habitats. In the face of

such limited sampling, predictive modeling of

habitat suitable for these taxa can be a useful

and cost‐effective tool that can help guide

research surveys and support conservation

action (Guinotte et al., this volume).

Two sets of predictive habitat suitability models

for deep‐sea coral taxa have been developed for

this region. Davies et al. (unpublished)

developed a predictive model for framework‐

forming stony corals in the region. Kinlan et al.

(2012) developed a set of models that included

the four major framework‐forming stony corals

(Oculina varicosa, Lophelia pertusa, Enallopsammia

profunda, and Madrepora spp.), as well as black

corals, gorgonians, soft corals, sea pens, and

stylasterid corals (Fig. 9). Both modeling teams

used the Maximum Entropy (Maxent; Phillips

et al. 2006) modeling approach relying on

presence‐only data.

Although a useful tool, these predictive habitat

models have not yet been fully validated by in‐

situ observations in the field. Additionally, the

model resolution (~ 370 – 1000 m) is generally

not fine enough to depict the mound features

that represent bioherms created by framework‐

forming stony corals. Maxent presence‐only

model results also tend to over‐predict suitable

deep‐sea coral habitat, both because of the

absence of high‐quality substrate data and in

cases where higher taxonomic levels (e.g.,

Order or Suborder) are modeled that contain

disparate numerous species with different

distributions. Nevertheless, the models closely

approximate the coral habitat zones identified

by Reed et al. (2013a, and Fig. 5) and suggest

that the region of potential coral habitat is

larger than was imagined a decade ago. The

Kinlan et al. (2012) model also accurately

predicted the occurrence of suitable O. varicosa

habitat north of the Oculina Bank HAPC (Fig.

9b). Surveys in 2010 confirmed the presence of

previously unknown Oculina mounds in this

area, providing vital information that led to the

expansion of the HAPC in 2015 (see below).

II.5. Other New Information from

Published Studies Since 2007  

Brooke et al. (2013) and Mienis et al. (2014)

noted that deep‐sea corals and associated fauna

occur in very dynamic and sometimes harsh

environments in this region, largely due to Gulf

Stream influences. Extreme and rapid changes

in bottom temperatures and other parameters

occur around Lophelia bioherms off Cape

Lookout, NC (Brooke et al. 2013, Mienis et al.

2014). Although temperatures can rise over the

expected lethal limit for Lophelia and persist at

those levels for almost a week, Brooke et al.

(2013) noted that L. pertusa colonies survived

these extremes. Mortality increased greatly

after a week at higher temperatures. While they

cautioned that the effects of temperature on

survival of L. pertusa were complex, and they

proposed that the species was unlikely to be

found in habitats where temperatures were

15°C or higher for extended periods.

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Figure 9. Maxent predictive habitat model for six different deepwater coral taxa in waters off the Southeast U.S.:

a) Lophelia pertusa; b) Oculina varicosa; c) Antipatharia (black corals); d) Gorgonian octocorals; e)

Pennatulacea (sea pens); f) Stylasteridae (lace corals). Warmer colors indicate greater likelihood of suitable

habitat, determined by a cross‐validation method that determined likelihood thresholds using the ratio of false

positive to false negative classification results when the model was tested on data left out of the fitting process.

STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Gartner et al. (2008) noted the occurrence of

large aggregations of mesopelagic fishes and

macronekton invertebrates in areas of the upper

continental slope off North Carolina. They

observed persistent aggregations near or on the

bottom that appeared to forego daily vertical

migrations. Aggregations were especially dense

in areas of steeply sloping ridges, canyons, or

other sharp relief, including Lophelia coral

mounds. Benthic fishes and invertebrates on the

coral mounds fed on species in the near‐bottom

aggregations, indicating that they clearly

contribute to energy flow around deep reefs.

Since their contributions might be substantial,

additional studies of midwater and benthic

interactions are needed.

Long‐lived deep‐sea corals continue to be

important proxies for understanding the

historical environment relevant to deep reefs

over the last 2000 years. Species of black corals

examined in paleoecological studies in the Gulf

of Mexico (e.g., Prouty et al. 2011, 2014) also

occur in the Southeast U.S. region. Sinclair et al.

(2011) estimated a bamboo coral (Keratoisis sp.)

from the region to be between 330 and 580

years old. These authors noted that the skeleton

of this species may provide a good record of

certain signals, but that more research was

needed to verify the utility of this coral.

II.6. Research Priorities

Ross and Nizinski (2007) identified a number of

deep‐sea coral research priorities for the region.

Also in 2007, the South Atlantic Fishery

Management Council (SAFMC) approved a

Deepwater Research and Monitoring Plan

(SAFMC 2007b) to guide conservation and

management of deep‐sea coral ecosystems in

the Southeast U.S. and to support mandates in

the Magnuson‐Stevens Fisheries Conservation

and Management Act (MSA). The plan has two

phases: 1) mapping and describing known and

expected deepwater coral ecosystems in the

region; and 2) determining the ecological role of

these ecosystems, especially as essential fish

habitat (EFH), and understanding the biology

and ecology of structure‐forming species. The

Deep Sea Coral Research and Technology

Program held research priority workshops in

2009 (NOAA 2010) and 2015 (Schull et al. 2016)

that built on and expanded research priorities

for the region (see below). Table 3 summarizes

research priorities identified in these efforts.

New priorities: While a number of the

priorities were addressed in the Deep Sea Coral

Research and Technology Program’s 2009‐11

research initiative, there remain significant gaps

in our understanding of these ecosystems,

including the following priorities.

Mapping and habitat characterization:

Mapping the location of deep‐sea coral and

sponge habitats remains a high priority.

High‐resolution multibeam sonar maps, ideally

with backscatter, are a foundation for most

other research, modeling, and conservation

(Correa 2012). This is especially true in this

region, where L. pertusa, E. profunda, and O.

varicosa, form coral mounds that can be

identified in multibeam surveys. The recent

discovery of hundreds of additional mound

features off Florida and Georgia suggest a

much richer deep‐reef province than previously

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Table 3. Summary of deep‐sea coral research priorities developed for the U.S. Southeast. NOAA held workshops

in 2009 (NOAA 2010) and 2015 (Schull et al. 2016). Specific recommendations have been paraphrased and

reorganized to highlight commonalities among specific recommendations.

Ross and Nizinski 2007 SAFMC 2007b NOAA 2010 Schull et al. 2016

Map: Detailed mapping of the Map: Map the distribution of Map: Inventory existing data; Map: What topographic features are

southeastern U.S. shelf edge and deepwater coral ecosystems in complete multibeam sonar associated with deep‐sea coral

slope (esp. multibeam in the the Southeastern U.S. EEZ; assess surveys for proposed deepwater ecosystems? Where are the hotspots

depth range of 350‐800 m). their condition Coral HAPCs, and explore and

characterize sites of interest

identified within the C‐HAPCs

and what is their spatial extent?

Biodiversity & Habitat

Associations: Document species

composition and distributions of

deep corals within the region;

document the whole living

habitat matrix and associated

fauna.

Biodiversity & Habitat

Associations: Describe and

inventory biota of deepwater

coral ecosystems; Describe the

population dynamics,

movements and habitat

associations of both economically

and ecologically important

associated species. Determine the

nature, patterns, and processes of

communities of microbial coral

associates.

Biodiversity & Habitat

Associations: Describe types,

strengths, and patterns of habitat

association of other species with

deep‐sea corals and sponges

Biodiversity & Habitat Associations:

Which species of deep‐sea corals and

sponges are present in these

ecosystems? What are the

relationships between deep‐sea corals

and sponges and their epifauna?

What drives community structure

differences among sites?

Population connectivity: Reproduction & Population Reproduction & Population Reproduction & Population

Genetic studies for the major Connectivity: Describe Connectivity: Understand the Connectivity: What is reproductive

coral species and dominant reproductive strategies biology and ecology of deep‐sea biology and ecology of deep‐sea

associated fauna to examine (gametogenic cycles, sex ratio, corals and Sponges; including: corals and sponges (larval biology,

taxonomic status, dispersal, fecundity, larval development demographics of deepwater coral colony size‐frequency distribution,

relationships among coral modes) of priority structure– and sponge species; recruitment, recruitment, environmental drivers,

banks, and community genetics forming groups, including

scleractinians (L. pertusa, E.

profunda, M. oculata), octocorals,

antipatharians and stylasterids.

Describe the genetic

characteristics of structure‐

forming coral populations.

connectivity and reproduction.

Conduct genetic analysis and

determine population dynamics;

understand the connectivity

between shallow, mesophotic,

and deep‐sea corals.

etc.)? How are populations connected

and what factors shape the genetic

connectivity of deep‐sea coral

ecosystems in the South Atlantic (e.g.

stepping stones, dispersal barriers)?

Food webs: Trophodynamics

studies of coral banks and

surrounding areas (whole water

column)

Food webs: Describe food web

dynamics of DWCEs

Food webs: What is trophic ecology

of deep‐sea coral and sponge

ecosystems? What are the food

sources?

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Ross and Nizinski 2007 SAFMC 2007b NOAA 2010 Schull et al. 2016

Age and growth: Understand the Age and growth: Describe

age and growth of the major patterns and processes of colony

structure forming corals (e.g., growth and mortality of

Lophelia, Madrepora, bamboo important structure‐forming

and black corals) and evaluate species, and determine how they

the age, history and present are affected by environmental

status (accreting, eroding) of the factors and stressors

underlying coral mounds.

Anthropogenic impacts: Long‐

term monitoring, research,

education, and enforcement of

deep‐sea coral MPAs.

Monitoring of bottom fisheries.

Anthropogenic impacts: Map

human activities that may impact

deepwater coral ecosystems;

Identify and quantify natural and

anthropogenic stressors

Anthropogenic impacts: Map

fishing and non‐fishing activities;

Identify and assess impact to

areas from fishing and non‐

fishing activities; monitor coral

recovery; determine appropriate

mitigation strategies.

Anthropogenic impacts: What

interactions occur between fishing

gear from the golden crab, royal red

shrimp, and wreckfish fisheries and

deep‐sea coral and sponge

ecosystems? How do deep‐sea coral

communities differ inside and

outside of protected areas? How can

we assess the health and resilience of

deep‐sea coral and sponge

ecosystems? How much can deep‐sea

fish and benthic communities be

impacted and still maintain

sustainable fisheries? Are there signs

of recruitment or recovery in areas

that have been impacted in the past?

Environmental archives:

paleoclimate or paleo‐

environmental studies of corals

Environmental archives: Can we

reconstruct paleoclimates using deep‐

sea corals and sponges?

Physical factors and habitat

modeling: Describe the

physiographic environment of

deepwater coral ecosystems;

Describe relationships of coral

ecosystem composition, structure

and distribution and abiotic and

biotic factors; develop models to

enable predictions of coral

ecosystem status and trends.

Habitat suitability modeling:

Develop and groundtruth

modeling tools to predict where

corals are likely to occur.

Physical factors and habitat

modeling: What topographic features

are associated with deep‐sea coral

ecosystems? How accurate are model

predictions of deep‐sea coral habitat,

particularly for structure‐forming

taxa? What is the most useful level of

spatial and taxonomic resolution for

models to guide cruise planning and

management?

Climate change impacts: What are

the environmental limitations (i.e.,

temperature, depth, dissolved

oxygen, aragonite saturation, pH,

etc.) of deep‐sea coral and sponges, as

well as their associated species, and

what is the variability in these

environmental parameters? How do

deep‐sea corals calcify under low

aragonite saturation states and low

oxygen levels?

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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imagined – perhaps among the world’s richest.

Likewise, multibeam surveys of portions of the

Stetson Banks (S.W. Ross, unpubl. data)

revealed many more mounds and rocky hard‐

bottoms than previously known. Both these

areas should be a high priority for new surveys.

The key second phase is to better classify, map,

and characterize the coral and sponge

assemblages and their associated biological

communities in relation to underlying geology

and oceanography.

Modeling: Species and habitat distribution

modeling is becoming a fundamental tool in

every U.S. region. Needed steps in the region

include ground‐truthing existing coral models,

developing new ensemble models for both

corals and sponges, and developing approaches

to model abundance and diversity (not merely

presence) of these taxa. Improving the

environmental and biological data inputs to

these models is also a critical activity.

Population connectivity: New genetic

techniques are providing tools to better

understand the connectivity of populations at

both fine and ocean basin scales (Morrison et

al., this volume). Next generation sequencing is

a promising new methodology to answer these

questions. These genetic tools can complement

and link to other studies and modeling of

current dynamics (e.g., Hilário et al. 2015).

Climate and ocean acidification: Since 2007,

the potential impacts of ocean acidification

and climate change have been identified as

a significant research need. The Fifth Assess‐

ment Report of the Intergovernmental Panel on

Climate Change (IPCC) highlighted the limited

information available on the risks of climate

change and ocean acidification to the deep‐sea.

The importance of improving this understand‐

ing given the size of the deep‐sea region is

imperative (Hoegh‐Gulberg et al. 2014).

A number of recent laboratory studies have

been conducted to determine responses of L.

pertusa to decreased aragonite saturations levels

associated with lower pH (e.g., Form and

Riebesell 2012, Lunden et al. 2014). These

studies have shown that this species is capable

of continued calcification even under reduced

aragonite saturation levels. However, the long‐

term consequences of ocean acidification for L.

pertusa and other reef‐forming species, as well

as the underlying coral frameworks that they

build, remain unclear. There is an extensive

unanalyzed database of ocean carbonate

chemistry for this region (Ross and Brooke,

unpubl. data) that indicates that several deep‐

sea coral species occur in areas of low aragonite

saturation, as reported in the Gulf of Mexico

(Lunden et al. 2014). These extensive data on

carbonate chemistry and nutrients would be

valuable as model inputs as well as basic

environmental descriptors. Given the important

role that reef‐forming deep‐sea

corals play in the region, understanding

carbonate chemistry is especially important. A

better understanding of the synergistic effects

of changing temperatures, oxygen

concentrations, and current regimes due to

climate change is needed, as well as how these

factors may interact with ocean acidification.

Sponges: This chapter is the first effort to

summarize our knowledge of the deep‐water

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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sponge fauna of the region. Understanding

these species and the habitats they form lags far

behind that of deep‐sea corals. There are critical

data gaps at every level, including basic

taxonomy, life history, and ecology. In addition,

sponges and their rich endosymbiotic

microorganisms rank among the most

important targets for bioprospecting (Hogg et

al. 2010).

Two new international consortia (Deep‐sea

Sponge Grounds EcosystemS of the North

Atlantic [SponGES] and A Trans‐Atlantic

assessment and deep‐water ecosystem‐based

spatial management plan for Europe [ATLAS])

have begun research on sponge grounds in the

northeastern Atlantic. They hope to quantify

the ecosystem services these habitats provide,

and to identify the threats they face from both

commercial activities (e.g., fishing) and climate

change. These projects were established with

support from the European Commission’s

Horizon 2020 program. The U.S. will partner

with these consortia to explore and identify

sponge grounds in the Southeast U.S.

III. Update on Management

of Deep-Sea Corals and

Sponges

III.1. New Information on Impacts

and Stressors

III.1.i – Fishing

Fishing continues to be among the most

important stressors to deep‐sea coral

ecosystems in the Southeast U.S. However,

beginning in 2010, the potential of fishing to

adversely impact these ecosystems was

drastically constrained by historic management

actions proposed by the South Atlantic Fishery

Management Council and implemented by

NMFS (see below). Trawl fisheries have the

potential to damage both corals and sponges in

the region (van Dolah et al. 1987). The principal

fisheries of concern in deeper waters have been

the localized royal red shrimp (Pleoticus

robustus) trawl fishery and the golden crab

(Chaceon fenneri) trap fishery. These are small

fisheries operating in a limited geographic

range, but both had the potential to severely

damage deep‐sea coral habitats. In shallower

water on the shelf, the rock shrimp (Sicyonia

brevirostris) fishery had been a significant

concern off parts of Florida, particularly to

Oculina reefs (Reed et al. 2007, 2008) prior to the

implementation of vessel monitoring

requirements for the fishery. Roth (2007)

conducted interviews with stakeholders and

concluded that inadequate surveillance and

enforcement resulted in continuing illegal

snapper‐grouper fishing in the Oculina Bank

Habitat Area of Particular Concern (HAPC).

Although potentially illegal incursions by

shrimp vessels into the Oculina Bank HAPC

have been reported (SAFMC 2015), regulations

and requirements for vessel monitoring

systems (VMS) have reduced impacts of fishing

gear on O. varicosa coral. Lesser impacts to

shelf‐edge coral and sponge habitats may also

occur from limited bottom‐set longline fishing

(prohibited south of 27°10′N and elsewhere in

depths less than 50 fathoms [91 m]).

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Analyses of golden crab and royal red shrimp

fisheries conducted in association with SAFMC

fishery management plan amendments

(SAFMC 2009, 2013) have provided a much

better understanding of the geographic

footprint of these fisheries. While neither of

these fisheries specifically target areas with

corals, geographic and bathymetric overlap

between the fisheries and Lophelia habitat

indicates that these bottom‐contact fisheries are

the most likely to interact with these habitats.

Golden crab occur on both soft‐bottom and

hard bottom‐habitats, including Lophelia

mounds, at depths between 205 ‐ 1,007 m

(Wenner and Barans 1990, Reed and Farrington

2010, Reed et al. 2017). The current golden crab

fishery operates off eastern Florida, where

fishers deploy strings of 20 to 50 traps that can

stretch up to 8 km (SAFMC 2009). Fishers target

soft sediment areas adjacent to deepwater coral

habitat, but strong currents can carry the traps

into coral habitats. It is likely that the greatest

potential damage to deep‐sea coral and sponge

habitats would occur during retrieval of the

trap lines, when the trap ground line is hooked

with a grapple towed behind the fishing vessel.

The sweep of the traps and lines as they are

lifted off the bottom has the potential to

damage areas of coral habitat and cause

sediment re‐suspension that can suffocate coral

polyps. Lost traps and line have also been

observed near coral habitat (S. Brooke, pers.

comm). It is a small fishery, with approximately

5 to 6 vessels landing golden crab in recent

years (SAFMC 2009) and 11 permits authorized

in 2016 (NMFS South East Regional Office).

Royal red shrimp are found on the continental

slope at depths from 180 – 730 m, with the

largest concentrations of shrimp occurring

primarily off northeast Florida at depths of

between 250 – 550 m (Anderson and Lindner

1995, SAFMC 2013). Based on the analysis of

VMS tracks, the primary royal red shrimp

fishing areas are from around 26.4 o N to 30o N

off eastern Florida, between about 250 – 400 m

depths. This range overlaps with the shallower

end of the Lophelia reef distribution. Although

the royal red shrimp is not a federally managed

species in the South Atlantic, regulations for the

Stetson‐Miami Terrace deepwater Coral Habitat

Area of Particular Concern (C‐HAPC) were

designed to prevent trawling impacts to

Lophelia coral habitats from the fishery (see

below).

In June 2008, NMFS implemented a mandatory

Federal observer program to characterize the

South Atlantic penaeid and rock shrimp

fisheries (Scott‐Denton et al. 2011). The

commercial rock shrimp fishery is focused off

eastern Florida, from approximately St.

Augustine to Vero Beach – primarily at 25 – 65

m depths. Fishing at these depths potentially

overlaps with Oculina reef habitat. Invertebrates

other than crustaceans represented 6% of the

rock shrimp fisheries catch (Scott‐Denton et al.

2011), but these data were not reported in

sufficient taxonomic specificity to determine the

extent to which corals or sponges were

represented in the bycatch.

III.1.ii – Other stressors

Oil and gas development: A potential future

stressor in the region may be oil and gas

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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development and associated risks of oil spills.

Studies from the 1970s and 1980s indicated that

certain areas off the Southeast U.S. may contain

significant oil and especially gas resource

potential (BOEM 2014). In 2015, the Bureau of

Ocean Energy Management (BOEM) published

a Draft Proposed Program (DPP) for 2017‐2022

Outer Continental Shelf Oil and Gas Leasing.

The DPP identified an option for a potential

lease sale in an area located at least 50 miles

offshore from the coasts of Virginia, North

Carolina, South Carolina, or Georgia. However,

in March 2016, the U.S. Department of the

Interior removed the U.S. Atlantic region from

oil and gas exploration from the current (2017‐

2022) five‐year plan (BOEM 2016). Thus, risks

from this activity in the near future are not

expected unless this decision is overturned. In

addition to potential threats from U.S. oil and

gas development, Cuba is starting to develop

capabilities to extract oil and gas resources in

the Florida Straits within their EEZ. Any spills

associated with such development could also

threaten U.S. deep‐sea coral and sponge

habitats downstream. As described in the Gulf

of Mexico chapter (Boland et al., this volume),

the Deepwater Horizon disaster demonstrated

the potential for an oil spill to damage deep‐sea

corals. Additional fossil fuel development

projects proposed for Southeast Florida and

that have included surveys for potential

impacts on benthic habitats include a liquid

natural gas (LNG) port facility (Messing et al.

2006a) and a natural gas pipeline between

Florida and the Bahamas (Messing 2006b).

Renewable energy: In the Southeast U.S., as in

other U.S. regions, there is increasing interest in

offshore ocean‐based renewable energy.

Offshore wind energy is the most mature

technology, and it has been successfully

implemented around the world, especially in

Europe. There are currently no facilities under

construction in the Southeast U.S., and when

implemented these facilities will likely be sited

predominantly nearshore in relatively shallow

waters – with little impact to deep‐sea coral and

sponge habitats.

Various entities have expressed interest in the

potential for developing marine current energy

or ocean thermal energy conversion (OTEC)

projects offshore of southeastern Florida

(Vinick et al. 2012). The exceptionally strong

currents associated with the Gulf Stream

flowing through the Florida Straits off the

southeastern Florida coast, represent the best

ocean current resource for renewable energy

development in the United States. While still in

early planning stages, both facility siting and

transmission lines to bring electricity onshore

have a potential to damage vulnerable benthic

habitats.

Submarine cable deployment: Southeast

Florida is a major hub for communications

cables to the Caribbean and Central and South

America. Although the footprint of cables is

much smaller than that of fisheries, their

deployment can result in adverse impacts to

sensitive habitats (Hourigan et al. 2007,

Messing et al. 2012). The State of Florida has

been a leader in developing incentives for

companies to route cables away from

environmentally‐sensitive habitats, including

deep‐sea coral habitats.

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Invasive species: There have been no systematic

surveys for invasive species in deeper waters of

the Southeast region. However, since 2007,

observations of increasing numbers of the

introduced Pacific lionfishes, Pterois volitans

and P. miles have raised concerns (Morris and

Whitfield 2009) in waters shallower than 150 m.

Large numbers of lionfish have been observed

down to 100 m in surveys throughout the

region, (Meister et al. 2005), with some

observations extending to 300 m (Grant

Gilmore pers. comm. cited in: Muñoz et al.

2011). Harter et al. (2015) recently documented

thousands of lionfish at depths of 50 to greater

than 100 m in protected hard‐bottom areas

along the Southeast U.S. from Florida to North

Carolina. While unlikely to damage corals and

sponges directly, predatory lionfish have the

potential to reduce forage fish biomass in the

associated communities (Muñoz et al. 2011). In

studies of mesophotic reefs in the Bahamas,

lionfish predation on herbivorous fish caused a

shift in benthic community structure, with an

increase in algae relative to corals and sponges

at depths to 61 m (Lesser and Slattery 2011).

Climate change and ocean acidification. Since

2007, significant new information has become

available on the potential impacts of climate

change and ocean acidification on deep‐sea

corals. The IPCC Fifth Assessment Report

concluded that while most ocean warming has

occurred in shallow waters, warming has also

been observed in ocean depths between 700 and

2000 m from 1957 to 2010 and likely involves a

significant anthropogenic signal (Hoegh‐

Guldberg et al. 2014). Rising atmospheric CO2

poses a risk to deep‐sea coral and sponge

communities through increasing temperature

(e.g., Brooke et al. 2013), decreasing O2 (Keeling

et al., 2010, Stramma et al. 2010 ), decreased

productivity of surface waters and export of

food to the deep‐sea (Jones et al. 2014), and

ocean acidification with associated changes in

carbonate chemistry that affect coral

calcification (Guinotte et al., 2006). Based on

laboratory and in situ experiments, Brooke et al.

(2013) concluded that the upper lethal

temperature limit for L. pertusa in the Southeast

U.S. is near 15o C. A number of upper‐slope

areas within the region may already be facing

temperatures in this range, at least for short

periods (Brooke et al. 2013, Mienis et al. 2014).

In a similar manner, climate‐driven expansion

of the oxygen minimum zone could adversely

affect coral growth. Lunden et al. (2014) found

in laboratory experiments that both high tem‐

peratures (> 14o C) and low oxygen concentra‐

tions (1.5 ml/l) resulted in mortality of L. pertusa

collected from the Gulf of Mexico. There is also

the potential for climate change to shift the

periods of peak primary productivity, which

provide food to the seafloor. Food delivery

drives deep‐sea species population dynamics,

reproduction and species composition.

Ocean acidification may be of particular

concern in the Southeast U.S. given that this

region holds the most extensive deepwater

coral bioherms in U.S. waters. Recent studies

suggest that living L. pertusa may be able to

continue calcification even in the face of

projected shoaling of the aragonite and calcite

saturation horizons (e.g., Form and Riebesell

2012). However, the three‐dimensional dead

coral framework that composes the mounds

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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may weaken and become more susceptible to

bioerosion in under‐saturated waters (Maier et

al., 2013, Hennige et al. 2015). Projections based

on IPCC emission scenarios indicate that most

living L. pertusa in the Southeast U.S. region is

likely to remain above the aragonite saturation

horizon for the remainder of this century (Jiang

et al. 2010).

III.2. New or Planned

Management Actions

III.2.i – Fisheries management

As described by Ross and Nizinski (2007), the

South Atlantic Fishery Management Council

(SAFMC) was an early leader in deep‐sea coral

habitat protection. The 1984 Oculina Bank

Habitat Area of Particular Concern was the

world’s first deep‐sea coral protected area.

Helies et al. (2011) recently conducted the first

survey of socioeconomic impacts of the Oculina

Bank HAPC based on surveys primarily

targeted at the recreational, charter and

commercial fisher communities in nearby Fort

Pierce, Florida. They reported different

perceptions of outcomes among the commercial

bottom‐fish long‐liners (overwhelmingly

negative), rock‐shrimpers (short‐term negative,

but long‐term positive) of the regulations

implemented between 1984 and 2000.

In 2010, NMFS approved the SAFMC’s

Comprehensive Ecosystem‐Based Amendment

1 (SAFMC 2009). This management action,

under development since 2001, established five

deepwater Coral Habitat Areas of Particular

Concern (C‐HAPCs). The C‐HAPCs protect a

total area of 62,717 km2 (24,215 mi2), off the

coasts of the Carolinas, Georgia, and eastern

and southern Florida, containing complex

deepwater coral habitats (Fig. 6). The Cape

Lookout and Cape Fear C‐HAPCs protect

known Lophelia habitats off North Carolina. The

Stetson Reefs, Savannah and East Florida

lithoherms, and Miami Terrace (Stetson‐Miami

Terrace) C‐HAPC stretches from South

Carolina to the Miami Terrace and until 2016

represented the largest marine protected area

(MPA) in the U.S. Atlantic. It includes the vast

majority of Lophelia and Enallopsammia habitats

identified by Reed et al. (2013a) as well as

habitat predicted to include these framework‐

forming corals (see above). The Pourtalès

Terrace C‐HAPC protects a significant portion

of this the Terrace and its unique coral habitats.

Within most areas of the C‐HAPCs, all fishing

vessels are prohibited from using all bottom‐

damaging gear, including bottom longlines,

trawls (bottom and mid‐water), dredges, pots

or traps, or the use of an anchor, anchor and

chain, or grapple and chain. “Allowable Golden

Crab Fishing Areas” and four “Shrimp Fishery

Access Areas” were included in the Stetson‐

Miami Terrace and Pourtalès Terrace C‐HAPCs

in order to accommodate these small‐scale

traditional fisheries in historic fishing grounds.

Within C‐HAPCs, possession of corals (with the

exception of octocorals off Florida) is also

prohibited. Reed et al. (2017) reported that at

least eight documented or potential Lophelia

coral mound habitat areas overlap with the

Allowable Golden Crab Fishing Areas, and

recommended that mangers should adjust the

boundaries to protect vulnerable habitat while

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

allowing benthic fisheries to continue operating

in areas of soft‐bottom habitat.

In 2013, the SAFMC proposed further

adjustments to the Oculina Bank HAPC and the

Stetson‐Miami Terrace C‐HAPC (SAFMC 2013,

Martin 2014) based on new findings from

research by NOAA’s Deep Sea Coral Research

and Technology Program as described above.

In addition, examination of museum records of

L. pertusa from off Cape Lookout, NC provided

a target location for additional multibeam sonar

mapping and the discovery of numerous low

profile mounds north of the Cape Lookout C‐

HAPC (Ross et al. 2012). A northward

expansion of the boundaries of this C‐HAPC to

include these mounds was also included in

Coral Amendment 8. Coral Ammendment 8

was approved by NMFS in 2015.

In addition to the C‐HAPCs, the SAFMC’s

Snapper‐Grouper Amendment 14 established a

series of deepwater marine protected areas

(MPAs) along the shelf edge (Fig. 6). NMFS

approved the amendment in 2009. The MPAs are

designed to protect seven long‐lived ʺdeepwaterʺ 

grouper and tilefish species (speckled hind,

snowy, yellowedge, Warsaw and misty groupers;

and golden and blueline tilefish), but also serve to

protect corals and sponges from fishing impacts.

The MPAs range in size from 27 km2 to 514 km2.

Fishing with potentially‐damaging bottom‐

contact gear, including bottom longlines, bottom‐

trawls, gill nets, golden crab trap or other traps is

prohibited, as well as fishing for or possession of

any snapper‐grouper species. Although much

smaller, the SAFMC has also approved a number

of Spawning Special Management Zones and

submitted these for approval by NMFS.

Together these actions have significantly

decreased the potential for damage from

fisheries to deepwater coral and sponge

habitats in the region.

III.2.ii – Other resource management

Oil and Gas: The Southeast U.S. Region

encompasses the Bureau of Ocean Energy

Management (BOEM) Planning Areas for the

Straits of Florida, South Atlantic, and portions

of the Mid‐Atlantic Outer Continental Shelf

(OCS). There has been renewed interest in oil

and gas exploration in the South Atlantic and

Mid‐Atlantic Planning Areas. At present, there

are no active oil and gas leases in any of these

areas, and no oil and gas lease sales are

proposed under the current Five Year Leasing

Program 2017‐2022.

Renewable Energy: BOEM is also responsible

for development of the Atlantic OCS wind,

wave, and other renewable energy resources.

As in other regions, there is increasing interest

in ocean‐based renewable energy development.

While wind energy utilization is expected to be

predominantly nearshore in relatively shallow

waters, the exceptional currents associated with

the Gulf Stream that flow through the Florida

Straits has catalyzed interest in hydrokinetic

energy development. The Florida Current is

close to the major metropolitan area of Miami,

and represents the best ocean current resource

for renewable energy development in the

United States. The proximity to relatively deep

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Figure 10. Major area‐based fishery management measures that protect deep‐sea coral and sponge habitats in the

South Atlantic Fishery Management Council region. These areas include Oculina Bank Habitat Area of

Particular Concern (HAPC) including the 2015 expansion; deepwater Coral HAPCs (2010 and expanded in

2015); and shelf‐edge Marine Protected Areas (MPAs).

STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

water also holds promise for potential ocean

thermal energy conversion (OTEC) projects.

While still in early planning stages, both facility

siting and transmission lines to bring electricity

onshore have a potential to damage vulnerable

benthic habitats, including deep‐sea coral and

sponge habitats on the Miami Terrace. Vinick et

al. (2012) assembled a seafloor survey

methodology and siting framework that

developers of marine and hydrokinetic projects

in this region could use. The authors identified

two major areas offshore southeastern Florida

that appear suitable for installing commercial

scale marine and hydrokinetic energy facilities,

including subsea electrical transmissions cables

to shore.

IV. Conclusions

New compilations and analysis of past

research, along with new benthic mapping and

research since 2006 have significantly improved

our understanding of the distribution of the

region’s deep‐sea corals and their associated

fauna.

 New mapping and surveys have confirmed

that the Southeast U.S. has by far the

nation’s highest concentration of

scleractinian deep‐sea coral mounds, and

probably ranks among the top such

provinces globally. Despite research

progress, much of this region remains

unexplored.

 Genetic studies of the important

framework‐forming species, Lophelia pertusa

and Oculina varicosa, have begun to reveal

information on population structure and

connectivity, while oceanographic studies

are elucidating factors that lead to coral

mound formation.

 In addition to branching stony corals,

gorgonians and black corals are important

contributors to habitat structure in areas of

hard bottoms. A more thorough cataloging

of these taxa has more than doubled the

number of species reported in the 2007

report.

 The first deep‐sea coral habitat suitability

models have been developed for the region,

but need to be validated using visual

surveys.

 Deep‐sea coral habitats are home to a large

number of associated species and act to

enhance local biodiversity.

The sponge fauna of the Southeast U.S. region

has not been systematically cataloged and

remains poorly known. Like corals, a number of

species form structurally complex habitats in

deeper waters, primarily on hard‐bottoms, and

likely contribute to habitat for numerous

species. There is a clear need for additional

research on the region’s sponges, including

their life histories and ecological roles.

The last decade has also seen the realization of

regional‐scale efforts to protect from fishing

impacts both deep‐sea coral reef provinces and

shelf‐edge coral and sponge habitats through a

series of deepwater Coral Habitat Areas of

Particular Concern (C‐HAPCs) and a network

of shallower MPAs, respectively. These new

fishing measures, developed by the South

Atlantic Fishery Management Council and

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

  

implemented by NOAA, have significantly

reduced the scope for fishing damage to deep‐

sea coral and sponge habitats.

V. Acknowledgements

We would like to thank Brian Kinlan and Matt

Poti for assistance on the predictive modeling

section. Stephen Cairns kindly reviewed the

taxonomy and coral distribution sections and

co‐authored the online annex. The chapter was

strengthened by reviews from Sandra Brooke,

Chip Collier, Martha Nizinski and Andrea

Quattrini. Robert McGuinn developed the coral

and sponge distribution and other maps for this

chapter, and Heather Coleman provided

editing assistance.

Online Annex – Comprehensive list of deep‐

sea corals in the U.S. Southeast region link:

https://deepseacoraldata.noaa.gov/library/2017‐

state‐of‐deep‐sea‐corals‐report

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Geographic Distribution 

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  

 Appendix 

Table A. Sites of major research expeditions (2007 – 2015) referenced in the chapter that included deep‐sea coral

and sponge research. (DSCRTP = NOAA Deep Sea Coral Research and Technology Program; OER = NOAA

Ocean Exploration and Research; HBOI = Harbor Branch Oceanographic Institution; NIOZ = Royal Netherlands

Institute for Sea Research; SEFSC = NMFS Southeast Fisheries Science Center; USF = University of South

Florida).

# Locations Year Vessel Description Program

1 Cape

Lookout

Lophelia

Banks ‐ off

North

Carolina

2009

2010

NOAA Ships

Nancy Foster

and Cape

Hatteras; R/V

Pelagia

6 month deployment of benthic Lander in Lophelia

habitats. PI: S.W. Ross

Oceanographic and geological studies, including box

and piston core samples

U. North

Carolina‐

Wilmington;

NIOZ

2 Off South

Carolina

2007 NOAA Ship

Nancy Foster

Multibeam and side‐scan sonar surveys to map bottom‐

topography on shelf edge and upper slope coral mounds

and other suspected areas of EFH on the outer

continental shelf and upper slope of the South Atlantic

Bight. PI: G. Sedberry

NOAA

Ocean

Exploration

3 Shelf‐edge

MPAs N.

Carolina to

Florida

2009 NOAA Ship

Pisces

ROV survey. PI: A. David NOAA

NMFS and

2010 M/V Spree ROV survey. PI: A. David CIOERT

2012 NOAA Ship

Pisces

ROV survey and multibeam mapping (261.0 km2 ). PI: S.

Harter

2013 NOAA Ship

Pisces

ROV survey and multibeam mapping (686.9 km2 ). PI: S.

Harter

2014 NOAA Ship

Nancy Foster

ROV survey and multibeam mapping (542.6 km2 ). PI: S.

Harter

2015 NOAA Ship

Pisces

ROV survey. PI: S. Harter

4 Off Georgia

and NE

Florida

2010 NOAA Ship

Pisces

ROV Surveys PI: G. Sedberry DSCRTP

2014 NOAA Ship

Okeanos

Explorer

Multibeam mapping of priority deepwater habitats OER

5 Jacksonville

Lithoherms,

Daytona

2011 NOAA Ship

Pisces

ROV survey and multibeam mapping (112 km2). PI: J.

Reed

DSCRTP

2010 NOAA Ship

Ronald H. Brown

ROV and multibeam surveys. 1575 km2 mapped. . PIs

SW Ross, S Brooke

DSCRTP

6 Off SE

Florida

2009 NOAA Ship

Nancy Foster

Multibeam mapping of priority deepwater habitats

within the Deepwater Coral HAPC. 580 km2 mapped. PI:

B. Costa

DSCRTP

2014 NOAA Ship

Okeanos

Explorer

Multibeam mapping of priority deepwater habitats OER

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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# Locations Year Vessel Description Program

7 Off SE

Florida,

Canaveral

Bioherms

2007 R/V Seward

Johnson

Fort Pierce and St. Lucie Pinnacles Submersible dives. PI:

J. Reed

HBOI

2009 R/V Seward

Johnson

Life on the Edge Expedition. 22 Submersible dives. PI: S.W.

Ross

http://fl.biology.usgs.gov/DISCOVRE/discovre_2009

index.html

DSCRTP,

CIOERT,

USGS

2010 NOAA Ship

Ron Brown

ROV and multibeam surveys. 1575 km2 mapped. PI:

S.W. Ross, S. Brooke

http://cioert.org/

DSCRTP

2011 NOAA Ship

Pisces

ROV surveys and some multibeam mapping (112 km2) DSCRTP

8 Oculina

Mounds,

Oculina

HAPC

2010 R/V Lost Coast

Explorer

Multibeam surveys – 7460 km2 mapped. PI: D. Naar DSCRTP,

USF

9 Off SE

Florida

Miami

Terrace and

Florida

Straits,

2007 R/V Seward

Johnson

Submersible dives. PIs: C Messing, J. Reed, S Brooke HBOI/OER

10 Florida

Straits,

Pourtalès

Terrace

2007 R/V Seward

Johnson

Submersible dives. PI: J. Reed HBOI/OER

2010 NOAA Ship

Ron Brown

ROV and multibeam surveys. PIs: S.W. Ross, S. Brooke DSCRTP

2011 NOAA Ship

Nancy Foster

ROV and multibeam surveys. 373 km2 mapped. PI John

Reed

DSCRTP

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Appendix – Deep-Sea Coral and Sponge Distribution Maps 

Map 1. Locations of deepwater (> 50 m deep) black corals (Order Antipatharia) recorded in the National Deep‐

Sea Coral and Sponge Database (as of November 2017).

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Map 2. Locations of deepwater azooxanthellate stony corals (Order Scleractinia) recorded in the National Deep‐

Sea Coral and Sponge Database (as of November 2017). Lophelia pertusa, Oculina varicosa, and

Enallopsammia spp. are the major framework forming species, with contributions from Madrepora spp.

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Map 3. Locations of deepwater non‐gorgonian alcyonacean corals (Order Alcyonacea) recorded in the National

Deep‐Sea Coral and Sponge Database (as of November 2017).

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Map 4. Locations of deepwater gorgonians (Order Alcyonacea in part [formerly Gorgonacea]) recorded in the

National Deep‐Sea Coral and Sponge Database (as of November 2017).

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Map 5. Locations of deepwater sea pens (Order Pennatulacea) recorded in the National Deep‐Sea Coral and

Sponge (as of November 2017).

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Map 6. Locations of deepwater lace corals (Family Stylasteridae) recorded in the National Deep‐Sea Coral and

Sponge Database (as of November 2017).

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STATE OF THE DEEP‐SEA CORAL AND SPONGE ECOSYSTEMS OF THE SOUTHEAST UNITED STATES

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Map 7. Locations of deepwater sponges recorded in the National Deep‐Sea Coral and Sponge Database (as of June

2016). Although sponges are widespread throughout the region, observations have not been systematically

recorded and organized, so records in the database are currently limited.

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